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Volume 38 1984 Number 1 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





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27 July 1984 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


LEE D. MILLER, President CHARLES V. COVELL, JR., 
KAROLIS BAGDONAS, Vice President Immediate Past President 
MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary 
J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


K. S. BROWN, JR. F. S. CHEW J. M. BURNS 
E. D. CASHATT G. J. HARJES F. W. PRESTON 
T. C. EMMEL E. H. METZLER N. E. STAMP 





The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229, U.S.A.; and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- 
temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- 
sion in the Sesiidae fascicle for the Moths of America North of Mexico, The dusky 
clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of 
willows and poplars. 


JoURNAL OF 
Tue LEPIDOPTERISTS’ SOCIETY 


Volume 88 1984 Number 1 


Journal of the Lepidopterists’ Society 
38(1), 1984, 1-12 


THE LIFE HISTORY AND ECOLOGY OF 
EUPHYDRYAS GILLETTII BARNES (NYMPHALIDAE) 


ERNEST H. WILLIAMS 


Department of Biology, Hamilton College, 
Clinton, New York 13323 


CHERYL E. HOLDREN AND PAUL R. EHRLICH 


Department of Biological Sciences, Stanford University, 
Stanford, California 94305 


ABSTRACT. Based on studies of several populations, the life stages of the montane 
butterfly Euphydryas gillettii and its natural history and ecology are described. E. gil- 
lettii shows unusual developmental flexibility in that it can diapause as second, third, or 
fourth instars, depending on climatic conditions; in addition, one population in a colder 
habitat is mostly biennial, while others are annual. In spite of this flexibility, the species 
has limited distribution in isolated populations over a narrow geographical range. 


Euphydryas gillettii Barnes occurs in the middle Rocky Mountains, 
ranging from western Wyoming, through northern Idaho and western 
Montana, and into Alberta (Ferris and Brown, 1981). While much work 
has been published on other species of Euphydryas in the past 20 years 
(Ehrlich et al., 1975; Cullenward et al., 1979; Brown and Ehrlich, 1980; 
Stamp, 1982), little has been known about E. gillettii. Until very re- 
cently (Williams, 1981; Holdren and Ehrlich, 1981), the only report in 
the literature on the biology of this species was that of Comstock (1940), 
which describes the eggs and early instars. 3 

We have studied E. gillettii in several locations recently and here 
report on its life history and ecology. Four populations have been 
observed extensively: natural populations in the Teton and Beartooth 
Mountains of Wyoming, and two populations introduced into Colorado 
from the Teton colony. In addition, several other populations have 
been visited. 


2, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Width of the head capsule, spine length, and body size for the different 
instars of Euphydryas gillettii. 





Length of spines 





Width of head a EO MI ETT 
Instar capsule (mm) Shaft (mm) Setae (mm) moving (mm) 
First 0.44 + 0.01 (30) 0.02 0.2-0.3 3-4 
Second 0.61 + 0.03 (81) 0.30 + 0.04 (28) 0.2-0.4 4-6 
Third 0.90 + 0.04 (46) 0.46 + 0.06 (43) 0.3-0.5 5-9 
Fourth 1.17 + 0.18 (84) 0.69 + 0.10 (39) 0.5—-0.7 9-13 
Fifth 1.47 + 0.18 (39) 0.74 + 0.10 (42) 0.6-0.9 12-18 
Sixth DADE = OMe2 (a) 0.74 + 0.16 (27) 0.7-1.2 15-30 





Study Sites 


The Beartooth population lives along a small stream in a montane 
meadow of 2620 m (8600 ft) elevation. The butterflies fly in an elon- 
gate area, roughly 60 m by 240 m, which is surrounded by coniferous 
forest of primarily Picea engelmanii. The highest density occurs in an 
area of secondary growth, where trees are scattered sparsely through 
a moist bottom area near the stream. 

The Teton population is the largest known for this species. The 
butterflies are widely scattered over an eastern facing slope at 2100 m 
(6900 ft) elevation, occurring in an area roughly 400 m by 1500 m of 
mostly herbaceous vegetation. Streams run down through this slope, 
and trees, mostly Picea and Populus tremuloides, grow along the stream 
beds. Adults are found throughout the slope. 

The two Colorado sites are in Gunnison County. One, adjacent to 
the Rocky Mountain Biological Laboratory at Gothic (2900 m, 9500 
ft), is similar to the Teton site. It consists of a moist meadow containing 
thick stands of willows on an east-facing slope, bounded by spruce 
forests, the East River, and the cliffs of Gothic Mountain. The second, 
Pioneer Resort (2700 m, 8800 ft), is less open than either the Gothic or 
Teton sites, but the flora is similar. 


Description of Life Stages 


Measurements of the head, spines, and body for the different instars are given in 
Table 1. 

Egg. Nearly spherical; rounded base with sides sloping in to flattened top. Approxi- 
mately 22 longitudinal ridges which extend most of distance down from apex, with 
irregular pitting on base; horizontal striations between ridges (Comstock, 1940). Color 
yellow-green when first oviposited (see Egg Development for color changes). Diameter 
0.78 + 0.02 mm (n = 11) and height 0.86 + 0.04 mm (n = 11) (eggmass shown in Fig. 
la). 

First Instar Larva. Head blackish brown with few thin, colorless setae. Body pale 
greenish yellow with colorless setae arising from 12 longitudinal rows of brown papillae. 
Appearance is of a pale body spotted with brown. Spiracles brown. Anal prolegs darker 
than other prolegs, which are concolorous with body; true legs brown (Fig. 1b). 


VOLUME 38, NUMBER 1 3 





Fic. 1. Life stages of Euphydryas gillettii: a, egg mass; b, first and second instars 
(prediapause); ec, third instars (postdiapause); d, fourth instars (postdiapause); e, sixth 
instar prepupa; f, pupa. 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Second Instar Larva. Head blackish brown with black setae. Body developing char- 
acteristic banding pattern of later instars: dorsal band pale yellow; dorsolateral band 
brown and irregular; lateral (stigmatal) band dull white; ventrolateral band light brown; 
and ventral band cream colored. Spiracles blackish brown. True legs brown. Crochets 
black; anal prolegs brown on outside. Branching spines develop from papillae and simple 
seta of first instar; shafts of spines light brown with black setae. Rows of spines as follows: 
one mid-dorsal in dorsal band; two in dorsolateral band, more dorsal row positioned 
caudal to second, and second on edge of next band; one row in lateral band; and two 
rows of small spines or tubercles adjacent to each other in ventrolateral band. Spines 
developed on all thoracic and abdominal segments, with exception of first and third rows, 
which are missing from thoracic segments (Fig. 1b). 

Third Instar Larva. Head capsule black with black setae. Body has same banding 
pattern of previous instar, but with deeper colors. Ventral band with thin mid-ventral 
brown line. Prolegs yellow with black crochets; anal prolegs dark brown on outside. 
Spiracles black. Shafts of spines blackish brown on all rows except mid-dorsal row, in 
which shafts are yellow-brown (Fig. lc). 

Fourth Instar Larva. Banding pattern further developed with greater contrast: dorsal 
band lemon yellow; dorsolateral band blackish brown with brown bases to spines; lateral 
band white with black spiracles, ventrolateral band brown; ventral band pale yellow with 
brown mid-ventral stripe. True legs black; prolegs yellow with brown bases and black 
crochets, and anal prolegs mostly black on outside. Shafts of all spines black, though with 
ring of lighter color at base of each, with yellow on light colored bands and brown on 
darker ones (Fig. 1d). 

Fifth Instar Larva. Colors and patterns as in previous instar, with following exceptions: 
dorsal stripe bright lemon yellow, dorsolateral band black, spines and setae jet black, and 
all prolegs yellow but dark on outside. 

Sixth Instar Larva. Continued development of previous banding pattern, with sharper 
contrast between bands. Midventral line blackish brown. 

Prepupa. Slight discoloration of last instar, with some shortening and thickening (Fig. 
le). 

Pupa. Ground color cream with black markings. Orange markings also occur except 
on wing cases; they are concentrated on abdominal segments, where there are seven 
orange warts per segment. Pupae average 16 mm long (Fig. 1f). 

Adult. Head and thorax black; abdomen black above and somewhat lighter under- 
neath. Palpi and legs concolorous with distinctive brownish orange color of postmedian 
band (this color is closest to the reddish orange of color 7B7 in Kornerup and Wanscher, 
1978; it is nearly identical to the orange-rufous, color II-11i, of Ridgway, 1912). Antennae 
black with thin white rings and with yellowish clubs. Dominant color of dorsal wing 
surface black; veins black; marginal band of orange and submarginal band of white 
much reduced, often disappearing in secondaries; postmedian band crossing both wings, 
3-4 mm wide, and prominent; median spot band white and reduced, disappearing by 
anal margin; discal cell of primaries with four alternating spots of white and characteristic 
orange-rufous color, with another spot of each color in postcellular space; secondaries 
with three spots of each color in cell and postcellular space; basal area black. Underwings 
with same patterning as above, but black color reduced and spots expanded; this is 
especially true on secondaries in median to basal area, where there is great expansion of 
orange-rufous color and where black is limited to borders of spots. Males smaller than 
females, with forewing length 16.5 to 23 mm (mean = 20.9, n = 162); for females, fore- 
wing length 20.0 to 25.5 mm (mean = 23.7, n = 199) (Fig. 2b). 


Ecology 


Oviposition. As reported by Comstock (1940), the larval host is Lo- 
nicera involucrata (Rich.) Banks (Caprifoliaceae), a shrub 0.5 to 3 m 
tall that grows in moist soil in thickets and wooded areas throughout 
the geographical range of E. gillettii and far beyond (e.g., California, 


VOLUME 38, NUMBER 1 5 


Mexico, Alaska, and Quebec). The leaves are glabrous, short-petiolate, 
elliptic-oblong to elliptic-obovate in shape, and 5-14 cm long and 2- 
8 cm wide (Hitchcock et al., 1959). Thus, the leaves are large enough 
to allow females to move completely to the underside of the leaves 
when ovipositing. Some authors (e.g., Tietz, 1972) have listed other 
larval foodplants, but eggs on or oviposition behavior near any plant 
other than L. involucrata is extremely rare. Of more than 600 egg 
masses seen in the Beartooth population, only four have been found 
on a plant other than L. involucrata; these occurred in 1982 on an 
unusually large and conspicuous specimen of Valeriane occidentalis 
Heller (Valerianaceae, a family related to the Caprifoliaceae). Post- 
diapause larvae may wander to other species of plants, however. 

Female E. gillettii oviposit mostly in late morning. Prior to ovipo- 
sition they fly slowly above the shrub and herbaceous layer, fluttering 
near or touching branches that are among the most apparent (highest 
or densest). They do not appear to follow vegetational edges. While 
searching for oviposition sites, they occasionally touch plants other than 
Lonicera, but then they usually fly on within 2 sec. 

Once a female does find L. involucrata, she flutters near the shrub, 
lands on a leaf, walks on it for a few seconds, and then flutters in the 
air, landing on the same or a different leaf. This process continues for 
1 to 80 minutes, and even when she is blown or chased from the shrub, 
she returns to the same leaf or to one quite near it. She gradually 
increases the time spent on one leaf, walking up and down the dorsal 
surface near the leaf midrib, repeatedly opening and closing her wings, 
and occasionally moving entirely to the underside of the leaf. After 
the female backs over or flips sideways to the underside of the leaf, 
there is an initial quiescent period of a minute or two which generally 
precedes oviposition. Sometimes she may return to the upper surface 
after remaining quiescent for a brief time, walk around the leaf again, 
and perhaps even move to another leaf. When she finally begins ovi- 
positing, she remains motionless with the wings usually held open (Fig. 
2b). Oviposition behavior of E. gillettii is quite similar to that described 
for E. phaeton (Stamp, 1982). Females appear to spend much time 
and effort assessing the potential oviposition site; individual females 
have been observed to spend more than two hours in the above be- 
haviors before actually beginning to oviposit. 

The leaves chosen for oviposition are always large and near the top 
of a growing stem. In the Beartooth population, 51% of the egg masses 
were on the highest leaf pair and 36% on the next highest leaf pair (2 
years, n = 453). Only one of 456 egg masses was found on the upper 
surface of a leaf, and only 7 of the 62 eggs from that egg mass hatched, 
while 30 were dislodged. The chosen leaves may or may not have other 


6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Euphydryas gillettii: a, prediapause feeding web, which becomes the hiber- 
naculum; b, ovipositing female; ec, parasitized fifth instar prior to emergence of the 
parasitoid. 


egg clusters already on them; in the Beartooth population, 44% (n = 
456) of all egg masses were on leaves that had another egg mass on 
the same leaf (23% of all leaves with eggs, n = 332), resulting in a 
mean of 1.87 clusters per leaf. Egg masses are also clumped in E. 
phaeton (Stamp, 1982). 

Approximately one-half of all egg clusters touch the leaf midrib. An 
ovipositing female faces the edge of the leaf and, while moving her 
abdomen back and forth, touches the lower leaf surface with the tip 
of the abdomen. If she then touches the midrib or another protruding 
leaf vein, she may use it as a guide in oviposition. Often she will use 
a previous egg mass as a guide. She lays the eggs row by row in both 
directions, and sometimes a second layer or more is oviposited upon 
the first. The far edge of the egg mass averages 2.0 cm from the edge 
of the leaf and the near edge 1.1 cm (n = 52), a distance which reflects 
the length of the body (roughly 1.6 cm). 

In the Beartooth population, egg clusters have ranged in size from 
23 to 310 eggs (n = 72), with a mean of 146 (Fig. la). In the Teton 


~] 


VOLUME 38, NUMBER 1 


population, the average size over a three year period was 130 eggs per 
mass (n = 189), while in Colorado the average was 128 eggs per mass. 
In contrast, the egg masses of E. editha contain 45 eggs on average 
(Labine, 1968), while those of E. phaeton contain 274 (Stamp, 1982). 
Oviposition in E. gillettii proceeds at an average of 3.8 eggs per minute 
(n = 48 clusters), requiring 38 min to lay an average sized cluster; E. 
editha oviposits at a slower rate, needing 30 min to produce its smaller 
cluster (Labine, 1968). Based on observations of 150 marked female E. 
gillettii seen to display pre-oviposition behaviors or to oviposit at least 
once, none oviposited more frequently than every other day. 

Egg Development. During the course of development in the Bear- 
tooth population, a mean of 13% of the eggs (n = 48 clusters) are lost 
from the egg mass due to dislodgement or detachment (19 eggs from 
a 146 egg average). Sometimes the edge of an egg mass peels away 
from a leaf, but most egg loss occurs where the eggs are more than 
one layer deep. The variance in egg loss per cluster is high, however, 
and most clusters lose few eggs. Presumably those eggs which detach 
from the leaves and fall to the moist, shady, predator-infested soil 
surface below do not hatch. 

In Colorado, up to 30% of the egg masses are lost entirely during 
the developmental period due to heavy predation. Furthermore, few 
egg masses escape without some predation; losses of roughly 10 to 20% 
of the eggs in a mass are common. The predators are the same for the 
eggs as they are for the larvae: erythraid mites, myrid bugs, beetle 
larvae, and browsing mammals, the latter including moose and cattle. 

The eggs change color during development from a pale straw-yellow 
when first oviposited, sometimes with a greenish tint, to a distinct gold, 
and then to darkening shades of red-brown. They become blue-gray 
about two days before hatching, a color which results from the for- 
mation of a dark head capsule beneath the white translucent egg shell. 

The eggs hatch from July into September, while the snow begins 
falling in late August in these mountainous areas. Eggs hatch in 23 to 
45 days in the Beartooth population, depending on the exposure of the 
oviposition site. In Colorado, the majority of the eggs hatch in 18-30 
days, although those masses that are produced late in the season de- 
velop more slowly. Eggs at the center of the egg mass are the first to 
hatch, and most eggs hatch within a two day time span (Williams, 
1981). A substantial fraction of the eggs, roughly 20% in the Beartooth 
population, hatch in early September after the leaves of L. involucrata 
have begun to wilt and turn yellow. 

Prediapause Larvae. Newly emerged larvae feed partially on the 
egg shells, and within 24 hours they migrate to the upper surface of 
the leaf, where they begin forming a communal feeding web (Fig. 1b). 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The oviposition leaf is the first feeding site and is the base of the web; 
it curls inwards and is bound ever more tightly as time passes. Predia- 
pause larvae feed only on the epidermis and parenchyma of the leaves, 
leaving behind the patterned network of veins. Feeding occurs during 
the day; nocturnal feeding has not been observed. Gradually more 
leaves are added to the feeding web by binding lower leaf pairs to the 
first leaf. In this way the communal web grows larger, sometimes with 
the incidental binding of grasses and other leaves that are adjacent to 
the hostplant leaves. The “knots” (Scudder, 1889) thus formed are 
quite apparent in the field since they generally occur at the apices of 
the most conspicuous stems (Fig. 2a). Because different egg masses are 
often oviposited on the same or adjacent leaves, the larvae in a single 
feeding web may be the products of several different egg masses, even 
when these egg masses hatch on different dates. 

Mortality is high during the prediapause period. Parasitic wasps 
identified as Benjaminia sp. (N. Stamp, pers. comm.) have been col- 
lected from the feeding webs, and the above-mentioned predators take 
a heavy toll. At least 80% of the larvae in the Beartooth population 
disappear before reaching winter diapause, while 50-60% of the Col- 
orado larvae die or disappear. 

Most, if not all, of the larvae that result from a single egg mass 
remain in the same feeding web overwinter. Though these hibernacula 
are well attached to the woody stems of the shrubs, most are dislodged 
by winter snow. 

Unlike the larvae of other well-studied Euphydryas, which diapause 
in the fourth instar, E. gillettii are apparently able to overwinter in 
response to environmental conditions as second, third, or fourth instars. 
For instance, the Beartooth colony, constrained by the rapid onset of 
winter at the end of the flight period, diapauses (first winter) in the 
second instar. In Colorado, where the two sites differ markedly in the 
length of both the larval and the food-plant growing season, the pop- 
ulations diapause at different instars even though they originated from 
the same parent colony in the Tetons. Like the original population, the 
larvae at 2440 m in Colorado reach the fourth instar, while at 2920 m 
they appear to overwinter successfully after the first molt but develop 
to the fourth instar given a sufficiently long summer (Holdren and 
Ehrlich, 1981). Overwintering larvae may pass through an extra molt 
before emergence, as occurs in E. editha (M. Singer, pers. comm.). 

Postdiapause Larvae. In Colorado and Wyoming the larvae termi- 
nate diapause soon after the snow melts, which in most years is late 
May at 2440 m (8000 ft) and mid-June at 2920 m (9600 ft). The larvae 
feed on newly formed buds of L. involucrata, boring holes into the 
larger apical buds and consuming entirely the smaller axillary buds 


VOLUME 38, NUMBER 1 9 


(Fig. lc). By the time the larvae have molted into the fifth instar in 
the annual populations, the leaves are slightly expanded, measuring 
roughly 2 cm in length. In postdiapause fourth instars in the biennial 
population, the larvae may still feed in aggregations (Fig. 1d) on rel- 
atively large and well developed leaves. Although many postdiapause 
larvae feed on shrubs bearing the previous year’s webs, like other Eu- 
phydryas species, some disperse. Extensive, characteristic feeding dam- 
age as well as postdiapause larvae have been observed on isolated L. 
involucrata shrubs on which there had been no prediapause larvae. In 
the Beartooth population, some postdiapause larvae have been found 
feeding on Castilleja linariaefolia Benth. (Scrophulariaceae), Valeri- 
ana occidentalis Heller (Valerianaceae) and Pedicularis bracteosa 
Benth. (Scrophulariaceae). All of these plants have iridoid glycosides, 
secondary compounds known from the host plants of other Euphydryas 
(Bowers, 1981). 

Diapause-related and postdiapause mortality appear to be quite high 
in Colorado. The number of postdiapause larvae found is consistently 
much smaller, by as much as two orders of magnitude, than the num- 
ber of large third instars observed shortly before diapause. Both post- 
diapause larvae and pupae may be parasitized, the latter in Colorado 
by the hymenopteran Ptermalus vanessae Howard, which oviposits 
into mature larvae or pupae. Parasitized larvae in the Beartooth pop- 
ulation cease feeding and movement in the fifth instar (Fig. 2c), and 
the Benjaminia parasitoid then emerges three to four weeks later. 

Most, if not all, larvae in the Beartooth population return to diapause 
for a second winter before pupating; they spend the first winter in the 
second instar and the second in the fifth instar. The second diapause 
apparently is not obligate, but the shortness of the growing season in 
this habitat has led at least part of the population into a two-year life 
span; Williams (1981) has demonstrated another adaptation in this pop- 
ulation for the cold climate, that of ovipositing so that the eggs are 
warmed maximally by the sun. A biennial life cycle has also been 
reported for Euphydryas maturna (Forster and Wohlfahrt, 1955), a 
close, European relative of E. gillettii. 

Larvae generally move away from the host shrubs for pupation (Figs. 
le & 1f), and the pupation sites are usually within 50 cm of the ground. 
While distinctive in color and pattern, the pupae are not easily found. 
Pupation requires about three weeks. 

Adults. The adults fly during a four week period from June to mid- 
August. As is typical for butterflies (Wiklund and Fagerstrom, 1977), 
males are the earliest to emerge and show the greatest wing wear early 
in the season, and the male to female ratio declines gradually through 
the flight season (Williams, in prep.). Males also fly earlier in the morn- 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ing than females and in relatively greater numbers on cloudy days. 
Males are much stronger fliers; though smaller (in accord with Singer, 
1982), they fly at faster speeds, are more difficult to catch, and are 
more difficult to manipulate when netted. 

These butterflies spend much of the day sunning near the ends of 
branches high in coniferous trees, typically with the wings open slightly 
more than 180 degrees. Males fly back and forth through the habitat 
more than females, while females fly down to nectar more frequently. 
Occasional individuals puddle in the afternoon when other activity is 
reduced. Nights are spent in trees at heights of at least 3 m. 

Mating is rarely observed because of the predilection of this species 
for the tops of nearby conifers. Chases of individuals near tree tops are 
common during the middle of the day, with males chasing both fe- 
males and other males. It remains curious, though, that males infre- 
quently chase females while females nectar in the herbaceous layer. 

The butterflies do not have to move far to nectar. There is a pro- 
fusion of flowers in the E. gillettii habitat, largely because it is moist, 
and they feed readily at the available blossoms. The commonest nectar 
source for the Wyoming populations is a white geranium, Geranium 
richardsonii Fischer and Trautvetter (Geraniaceae), which is also used 
in Colorado where the most important source is probably Erigeron 
peregrinus (Pursh) Greene (Compositae). After senescence of the pri- 
mary nectar source, E. gillettii in Wyoming turns readily to yellow 
composites, mostly several tall Senecio which begin blooming as the 
Geranium cease. Given the abundance of flowers and the relatively 
limited time spent nectaring, adult food resources would not seem to 
be a major limiting factor in the population dynamics of this species. 


DISCUSSION 


Euphydryas gillettii was originally described and placed in the ge- 
nus Melitaea by Barnes (1897) from material collected in Yellowstone 
National Park, Wyoming; M. glacialis (Skinner, 1921) is a synonym. 
Gunder (1929), in his reorganization of North American Euphydryas, 
recognized the relationship of E. gillettii to the other Euphydryas 
species and pointed out that it is likely the most primitive of the North 
American species. L. G. Higgins (1978) then revised the genus Euphy- 
dryas and placed E. gillettii in a new genus, Hypodryas, along with 
the Palearctic species E. maturna, E. intermedia, E. eduna, and E. 
cynthia. Phenetically, E. gillettii seems most closely related with those 
species, although comparison of early stages and allozyme frequencies 
would clearly be desirable. 

Following good taxonomic practice we have not accepted Hypodry- 
as as a genus; obligatory categories—genera, families, etc.—should be 


VOLUME 38, NUMBER 1 ll 


kept conservative to facilitate communication (Ehrlich and Murphy, 
1982). Hypodryas could be considered as synonymous with “the ma- 
turna species group’ or, at most, a subgenus. Euphydryas is a phenet- 
ically quite uniform group. Because the genus is now so widely dis- 
cussed in the non-lepidopterological literature, we would not suggest 
any change in the widely accepted generic name. 

Of current interest in the study of butterflies is whether or not the 
prior presence of eggs influences where a female lays her eggs. In 
several species—Battus philenor (Rausher, 1979), Pieris brassicae 
(Rothschild and Schoonhoven, 1977), and Anthocharis sara (Shapiro, 
1980)—active egg load assessment is indicated, and in all of these cases 
females avoid ovipositing where eggs currently are or recently have 
been. Female E. gillettii rarely avoid leaves that already have eggs; 
moreover, the egg clusters are grouped together more than one would 
expect if they were distributed in the environment at random (Wil- 
liams, 1981). The same is apparently true of E. phaeton (Stamp, 1982). 
Though there has been no previous support for positive egg load as- 
sessment, the grouping of eggs or egg clusters together may further 
enhance survivorship of larvae if there is a selective reason, such as 
predator avoidance or thermoregulation, for grouping the eggs togeth- 
er initially. Stamp (1981, 1982) has considered reasons for such a group- 
ing, though in her experiments, E. phaeton suffered increased parasit- 
ism when the groupings were too large. Because the larvae from 
different clusters of E. gillettii eggs do mix freely in communal feeding 
webs, the contagious distribution of clusters may be adaptive. 

E. gillettii displays sedentary behavior and occurs in localized col- 
onies with few populations known; these characteristics, along with the 
ease with which individuals may be caught, indicate that it could easily 
suffer from excessive human impact. How threatened the species may 
be is unknown, largely because it occurs in undisturbed mountain hab- 
itat, but much reduction in numbers in any one place could lead to 
the extinction of local colonies. Those who find a population in the 
field should exercise discretion when collecting, especially with fe- 
males. 

ACKNOWLEDGMENTS 


We thank Deane Bowers, Art Shapiro, and an anonymous reviewer for commenting 
on the manuscript. EHW was supported by grants from the Theodore Roosevelt Me- 
morial Fund of the American Museum of Natural History and from Wellesley College. 
The work of CEH and PRE was supported by a series of grants from the National Science 
Foundation, the most recent of which was DEB-8206961, and by a grant from the Koret 
Foundation of San Francisco. We thank the Brachman-Hoffman Foundation for support 
of publication. 


LITERATURE CITED 


BARNES, W. 1897. Some new species and varieties of Lepidoptera from the western 
U.S. Canad. Entomol. 29:39-42. 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Bowers, M. D. 1981. Unpalatability as a defense strategy of western checkerspot but- 
terflies (Euphydryas Scudder, Nymphalidae). Evolution 35:367-375. 

Brown, I. L. & P. R. EHRLICH. 1980. Population biology of the checkerspot butterfly, 
Euphydryas chalcedona. Structure of the Jasper Ridge colony. Oecologia 47:239- 
251. 

Comstock, J. A. 1940. Notes on the early stages of Euphydryas gillettii Barnes. Bull. 
S. Calif. Acad. Sci. 39:111-118. 

CULLENWABRD, M. J., P. R. EHRLICH, R. R. WHITE & C. E. HOLDREN. 1979. The 
ecology and population genetics of an alpine checkerspot butterfly, Euphydryas 
anicia. Oecologia 38:1-12. 

EHRLICH, P. R. & D. D. MurpHy. 1982. Butterfly nomenclature: A critique. J. Res. 
Lepid. 20:1-11. 

EHRLICH, P. R., R. R. WHITE, M. C. SINGER, S. W. MCKECHNIE & L. E. GILBERT. 1975. 
Checkerspot butterflies: A historical perspective. Science 188:221-228. 

FERRIS, C. D. & F. M. BROWN. 1981. Butterflies of the Rocky Mountain States. Univ. 
Oklahoma Press, Norman. 

Forster, W. & T. A. WOHLFAHRT. 1955. Die Schmetterlinge Mitteleuropas. Franck- 
h’sche Verlagshandlung, Stuttgart. 

GUNDER, J. D. 1929. The genus Euphydryas Scud. of boreal America (Lepidoptera, 
Nymphalidae). Pan-Pac. Entomol. 6:1-8. 

HiccIins, L. G. 1978. A revision of the genus Euphydryas Scudder (Lepidoptera: Nym- 
phalidae). Entomol. Gaz. 29:109-115. 

HitcHcock, C. L., A. CRONQUIST, M. OWNBEY & J. W. THOMPSON. 1959. Vascular 
Plants of the Pacific Northwest. Part 4. Ericaceae Through Campanulaceae. Univ. 
Washington Press, Seattle. 

HOLDREN, C. E. & P. R. EHRLICH. 1981. Long range dispersal in checkerspot butter- 
flies: Transplant experiments with Euphydryas gillettii. Oecologia 50:125-129. 
KORNERUP, A. & J. H. WANSCHER. 1978. Methuen Handbook of Colour. 3rd ed. Meth- 

uen, New York. 

LABINE, P. A. 1968. The population biology of the butterfly, Euphydryas editha. VIII. 
Oviposition and its relation to patterns of oviposition in other butterflies. Evolution 
22:799-805. 

RAUSHER, M. D. 1979. Egg recognition: Its advantages to a butterfly. Anim. Behav. 27: 
1034-1040. 

RipGway, R. 1912. Color Standards and Color Nomenclature. Publ. by the author; 
Washington, D.C. 

ROTHSCHILD, M. & L. M. SCHOONHOVEN. 1977. Assessment of egg load by Pieris 
brassicae (Lepidoptera, Pieridae). Nature 266:352-355. 

SCUDDER, S. 1889. The Butterflies of the Eastern United States and Canada. W. H. 
Wheeler, Cambridge. 

SHAPIRO, A. M. 1980. Egg-load assessment and carryover diapause in Anthocharis 
(Pieridae). J. Lepid. Soc. 34:307-315. 

SINGER, M. C. 1982. Sexual selection for small size in male butterflies. Am. Nat. 119: 
440-443. 

STAMP, N. E. 1981. Effect of group size on parasitism in a natural population_of the 
Baltimore checkerspot Euphydryas phaeton. Oecologia 49:201-—206. 

STAMP. N. E. 1982. Selection of ovipositon sites by the Baltimore Checkerspot, Euphy- 
dryas phaeton (Nymphalidae). J. Lepid. Soc. 36:290-302. 

TiETZ, H. M. 1972. An index to the described life histories, early stages and hosts of 
the macrolepidoptera of the continental United States and Canada. Vol. I. Allyn 
Museum of Entomology, Sarasota, Fla. 

WIKLUND, C. & T. FAGERSTROM. 1977. Why do males emerge before females? Oeco- 
logia 31:153-158. 

WiLuiAMs, E. H. 1981. Thermal influences on oviposition in the montane butterfly 
Euphydryas gillettii. Oecologia 50:342-346. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 13-14 


CORRECT NAME FOR THE NEOTROPICAL SQUASH-VINE 
BORER (SESIIDAE: MELITTIA) 


VITOR O. BECKER! 
AND 
THOMAS D. EICHLIN? 


ABSTRACT. The identity of the species of squash-vine borer occurring in Central 
and South America on cultivated Cucurbitaceae is established as Melittia pulchripes, not 
M. satyriniformis, which is a junior synonym of the Eastern squash-vine borer, M. 
cucurbitae. A lectotype is designated for M. riograndensis, a name which is then syn- 
onymized under M. pulchripes. 


For more than a century the Melittia species whose larvae are com- 
monly found boring in stems of many cultivated species of Cucurbi- 
taceae in Central and South America has been referred to in the lit- 
erature as M. satyriniformis Hiibner. In a study by Duckworth and 
Eichlin (1978) it was found that in the Western Hemisphere these 
borers belong to a complex of three closely related species: cucurbitae 
(Harris), satyriniformis Hiibner, and a third which they described and 
named calabaza. According to these authors (1973:154), the three species 
of the complex are easily distinguished by their external features and 
genitalia. A fourth species, pauper LeCerf, apparently occurs only in 
the vicinity of Lima, Peru. Both cucurbitae and calabaza are restricted 
to the United States and Mexico and are sympatric in the southern part 
of their range. The species distributed from Guatemala through Cen- 
tral and South America was regarded by them as satyriniformis, fol- 
lowing the use of earlier authors. 

Heppner and Duckworth (1981:26) established that satyriniformis 
is a junior synonym of cucurbitae. This was based mainly on the fact 
that Hiibner stated that the type locality of satyriniformis was “Geor- 
gia” and therefore, must be conspecific with cucurbitae, the only squash- 
vine borer from the region. 

We have examined the syntypes of riograndensis Bréthes (1920:284) 
and found that they are the same species as the Central and South 
American species previously and currently misnamed as satyrinifor- 
mis. Two male syntypes of riograndensis were located in the Museo 
Argentino de Ciencias Naturalis (MACN), Buenos Aires, both bearing 
identical labels in Bréthes’ hand-writing: “17”; “E. Ronna, vi. 1919, 


>> “<e >> < 


Pelotas’’; ““Type’’; ‘“Melittia riograndensis Breth.”’; (red rectangle). The 


‘Centro de Pesquisa Agropecuaria dos Cerrados, P.O. Box 70-0023, 73300-Planaltina, DF, Brazil. 
? Division of Plant Industry, Insect Taxonomy Laboratory, California Department of Food and Agriculture, Sacra- 
mento 95814, USA. 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


male specimen which had been dissected is here designated as the 
lectotype; the second male becomes a paralectotype. These two spec- 
imens are covered by mold, and their external features are somewhat 
obscured; however, the genitalia are identical to those of satyriniformis 
(sensu authors, including Duckworth and Eichlin, 19738:fig. 3c), bear- 
ing the peculiar quadrate expanded process at the center of the valva. 
They also agree with the genitalia of a specimen reared by the senior 
author from stems of Cucurbita sp. at Turrialba, Costa Rica. 
However, the oldest available name for this species is Melittia pul- 
chripes Walker (1856:67). Syntypes in the British Museum (Natural 
History) were examined and a lectotype designated (Duckworth and 
Eichlin, 1978:21). At this time it was determined to be conspecific with 
the Central and South American squash-vine borer. The previous ref- 
erences to satyriniformis for the Neotropical squash-vine borer should 
in fact be applied to pulchripes. Also, riograndensis now becomes a 
synonym of pulchripes (NEW SYNONYMY). 
The following is a summary of the species comprising the squash- 
vine borer complex: 
Melittia cucurbitae (Harris)—eastern half of United States, Gulf 
Coastal areas of Texas and Mexico to near Guatemala. 
Melittia calabaza Duckworth and Ejichlin—Arizona, central and 
western Texas, interior areas of Mexico to west coast. 
Melittia pulchripes Walker—Guatemala south throughout Central 
and South America to southern Brazil. 
Melittia pauper LeCerf—currently recorded only from Peru. 


LITERATURE CITED 


BRETHES, J. 1920. Insectos utiles y daninos del Rio Grande do Sul y de La Plata. Anales 
de la Sociedad Rural Argentiana 54:281-290. 
DuCKworTH, W. D. & T. D. EICHLIN. 1978. New species of clearwing moths (Lepi- 
doptera: Sesiidae) from North America. Proc. Entomol. Soc. Wash. 75:150-159. 
1978. The type-material of Central and South American clearwing moths (Lep- 
idoptera: Sesiidae). Smithson. Contr. Zool. 261:1-28. 

HEPPNER, J. B. & W. D. DuCKworTH. 1981. Classification of the superfamily Sesioidea 
(Lepidoptera: Ditrysia). Smithson. Contr. Zool. 314:1-144. 

WALKER, F. 1856. List of the Specimens of Lepidopterous Insects in the British Mu- 
seum. Part 8, 271 pages. London: British Museum. 





Journal of the Lepidopterists’ Society 
88(1), 1984, 15-22 


LIFE HISTORIES OF FOUR SPECIES OF 
PHILIRIS ROBER (LEPIDOPTERA: LYCAENIDAE) 
FROM PAPUA NEW GUINEA 


MICHAEL PARSONS 


Insect Farming and Trading Agency, Division of Wildlife, 
P.O. Box 129, Bulolo, Morobe Province, Papua New Guinea 


ABSTRACT. The life histories of four species of Philiris, P. helena Snellen, P. agatha 
Grose-Smith, P. intensa Butler and P. ziska Grose-Smith, together with notes on their 
biologies, are described and illustrated. 


Life histories of species of the genus Philiris Rober (Lycaenidae) 
from the Melanesian region have been little studied, but Forbes (1977) 
has recently detailed the life history of P. moira Grose-Smith from 
Papua New Guinea. Common and Waterhouse (1981) have briefly 
outlined the life history of P. innotata Miskin from Australia. 

In the past, difficulty has been experienced with the placing of the 
correct females with the males of certain species of Philiris, but Sands 
(1979, 1980, 1981) has done much towards clarifying the taxonomy of 
the genus. The phyletic arrangement of all species, however, will re- 
main unclear until the biologies of further species are known. It is 
hoped that the addition to the literature of four new Philiris life his- 
tories, and information about the morphology of their early stages, will 
assist in such a study. I am also preparing to describe the life histories 
of Philiris harterti Grose-Smith, P. diana Waterhouse & Lyell, P. vi- 
oletta Rober, and P. praeclara Tite from Papua New Guinea, all of 
which feed on Litsea (Lauraceae). 

Figs. 4 and 5 of the mature larva and pupa of P. moira are included 
here for ease of comparison. All life histories were studied from the 
Bulolo valley in the Morobe Province from approximately the center 
of the 10 km grid square reference DN50O. The early stages of all 
species show remarkable camouflage against their foodplants. The du- 
ration of the life cycle of each species was about one month from egg 
to adult. 


Philiris helena Snellen 


Egg. Diameter 0.75 mm; white, spherical when viewed from above, oval from the 
side, ventrally flattened; micropylar pit surrounded by six smaller pits; egg honeycombed 
with larger, regular, ovoid pits, the walls of which are produced into long, outward 
curving spicules which shorten gradually towards median line. 

Larva. First instar. 1.55 mm in length, 0.5 mm in width; oval and elongate in dorsal 
profile; head pale brown; body pale green, edged with pale yellow; hirsute, fringed with 
fine white setae. 

Second instar. 3.5 mm by 1 mm; anterior end slightly wider; head pale brown; body 
pale green with white middorsal line, broken at each segment. 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Third instar. 5 mm by 2 mm; similar to second but white middorsal line bordered 
with tan brown. 

Fourth instar. 10 mm by 4.5 mm; similar to third but middorsal line pinkish brown 
pattern encircled by cream; body color pink where larva has been feeding on small, 
young, pink leaves of foodplant, otherwise pale green; setal fringe 1.5 mm in length. 

Fifth instar. 16 mm by 6.5 mm; similar to fourth but middorsal line broader (3 mm), 
forming a diffuse pattern of four lines of white dashes; laterally this pattern is continued 
but is fainter (Figs. 2 and 6). 

Pupa. 11 mm in length, 5 mm in width; oval from above, ventrally flattened; hirsute, 
covered by fine pubescence of soft, white setae; dimorphic depending on substrate color 
being either pale green or pale brown; abdomen greenish yellow dorsally with two pale 
green dorsolateral lines either side of wider green middorsal line which continues onto 
thoracic segments. Supported by cremaster and fine silk girdle (Fig. 7). Duration, 12 
days. 


BIOLOGY. The foodplants are Macaranga aleuritoides F. Muell. and 
M. quadriglandulosa Warb. (Euphorbiaceae). Both species can grow 
up to 8 m in height. The leaves (Fig. 3) of both are broad and dark 
green. Those of M. quadriglandulosa are peltate, at the base (up to 35 
mm in diameter) with a pointed tip and a serrated edge. Those of M. 
aleuritoides have five lobes are semipalmate and often grow to 70 mm 
in diameter. Both possess four to eight shiny, red, ovate glands on the 
upperside of the leaf bases which appear to be attractive to various. 
species of ants. Gressitt and Nadkarni (1978, p. 114) mention that these 
extrafloral nectaries are also attractive to several families of flies. M. 
quadriglandulosa has a coarse felt-like covering of hairs on its leaves. 
The sap of both is sticky, clear and gelatinous. The foodplants are 
common throughout the Lae-Wau region in regrowth areas from sea 
level to 1200 m. 

Adults of P. helena can often be seen in abundance where the food- 
plant grows. They rest on the upper surface of the leaves and may be 
frequently seen drinking on damp sand. 

Eggs are usually laid single on the leaf petiole and adhere to the 
long, felty hairs (Fig. 1). The larvae when young eat only the lower 
epidermis, creating windows in the leaf. Later they eat many full holes 
(Fig. 3), skeletonizing the leaf with no more than the veins to hold its 
former shape. Ants are almost always present on the Macaranga food- 
plants, but their association with the P. helena larvae is minimal, and 
they seem more preoccupied with the plant glands. 

Pupae are always attached to the old leaf bracts at the base of the 
main stem. If the bracts are dry and brown, then the pupae tend to 
be brown also. If the bracts are still pale green, then the papae are of 
the same color. 

From a final instar larva of P. helena which was collected wild, a 
small tachinid fly emerged. The species appears to be prone to attack 
by these parasitoids because numerous mummified skins of final instar 


VOLUME 38, NUMBER 1 17 





Fics. 1-3. Philiris early stages and feeding damage: 1, egg of P. helena on petiole 
of Macaranga aleuritoides; 2, eggs (above bract) and mature larva and empty pupal 
case of P. helena on bract of M. aleuritoides; 3, damage to leaves of a) M. aleuritoides, 
b) M. quadriglandulosa, c) M. involucrata, d) Ficus calopilina (by P. moira). 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


larvae have been observed on the foodplants, each of which bore the 
ventral exit hole of a fly larva. 


Philiris agatha Grose-Smith 


Egg. Like that of P. helena. 

Larva. The first three instars of P. agatha closely resemble those of P. helena but 
fourth instar pattern discernibly different. Larva of P. agatha slightly slimmer and more 
elongate than P. helena. 

Fourth instar. 11] mm by 4 mm; head tan brown; body pale olive green; middorsal 
pattern bright white commencing from behind prothorax and composed of three parallel 
dashes on each segment which converge to form single broad line at anal end of larva; 
laterally, larva patterned with narrower parallel white lines; setal fringe 1.5 mm in length. 

Fifth instar. 18 mm by 6 mm; similar to fourth (Fig. 8). 

Pupa. 12 mm in length, 5 mm in width; hirsute, pale olive green, boldly patterned 
with white; prothorax with white, triangular markings above eyes; mesothorax with two 
broad white lines which curve around middorsal line; middorsal abdominal pattern com- 
posed of broad white dashes either side of midline; laterally abdomen bears single, broad, 
white, wavy line. Supported by cremaster and fine silk girdle (Fig. 9). Duration, 12 days. 


BIoLoGy. The foodplant is Macaranga involucrata (Roxb.) Bail. 
(Euphorbiaceae), a tree which grows to 7 m in height. The leaves (Fig. 
3) are variable in shape. They are either rounded or have three or five 
lobes, the central lobe being the longest. The leaves are covered with 
a fine white pubescence which makes them extremely soft and felt- 
like to the touch. They bear four small, often vestigial, glands on the 
leaf upperside near the petiole. The sap is clear, slightly sticky, and 
can have a strong camphor-like smell. It is a common plant of regrowth 
areas around the Bulolo valley. 

Adults of P. agatha are less frequently seen than those of P. helena 
and may be classed as occasional in their habitat. They sometimes drink 
on damp sand and mud at creek margins and are very fast flying. 

Eggs are usually laid on the underside of a small new leaf, or its 
petiole, at the apex of the foodplant. The larvae commence feeding on 
the young leaves. Then, as they grow, they move to feed on lower, 
older leaves. Leaf damage is shown in Fig. 3. No actual attendance of 
the larvae by ants was noted, although there were often brown tree 
ants on the foodplant. 

The monomorphic pupae are invariably attached to the underside 
of a very young leaf (hardly larger than the pupa) at the apex of the 
foodplant. They match well the felty appearance and color of these 
leaves. 


Philiris intensa Butler 


Egg. Diameter 0.5 mm; white; hemispherical, with a regular covering of spicules. 

Larva. First instar. 0.75 mm in length, 0.25 mm in width; oval in shape; head tan 
brown, lying well beneath prothorax; uniform greenish yellow; fringed with fine white 
setae. 


VOLUME 38, NUMBER 1 19 


Second instar. 3 mm by 1.5 mm; similar to first but with middorsal line patterned 
with reddish brown spots with dark green centers. 

Third instar. 6 mm by 2.5 mm; similar to second but brown middorsal line broken 
centrally by two white spots. 

Fourth instar. 8.5 mm by 3.5 mm; similar to third but spiracles white. 

Fifth instar. 10 mm by 6.5 mm; similar to fourth but ground color darker green and 
matches that of leaf on which larva feeds; middorsal line of brown spots with white 
centers; setae fringe 1 mm in length, armed with prominent, outwardly directed barbs 
arranged in alternating rows along each hair (Fig. 10). 

Pupa. 9 mm in length, 5 mm in width; smooth, not hirsute; apex of mesothorax with 
diffuse pattern of brown and white, ringed by olive green to edge ot pale yellow wings; 
abdomen pale lime-green with brown and white middorsal pattern on segments 1-5; 
spiracles white, those on first abdominal segment encircled by brown spots. Supported 
by cremaster and fine silk girdle (Fig. 11). Duration 10 days. 


BIoLocy. The foodplant is Pipturus argenteus Willd. (Urticaceae), 
a common plant of creekside and regrowth areas, which grows to about 
5 m tall. The leaves are pale to dark green and ovate with pointed tips 
and serrated edges. They average about 15-20 cm long and are felty 
to the touch. They are covered with minute white hairs. The small 
clusters of rounded, dimpled, opaque white fruit are gelatinous and 
are borne in alternating rows along fruit stalks. 

Adults of P. intensa are commonly seen near the foodplant, and 
males are especially fond of drinking on damp sand. 

Eggs are laid singly on the leaf underside, usually near the base. At 
all stages larvae eat the upper epidermis of the leaf and leave a char- 
acteristic long, narrow feeding trail of a meshwork of small veins. On 
occasion they were attended by small brown ants. 

Pupation is always on the upperside of a large or small leaf of the 
foodplant and along the main vein just before it joins the petiole. 

A small (8 mm long) orange ichneumonid wasp parasitoid was reared 
from a wild collected pupa. 


Philiris ziska Grose-Smith 


Egg. Diameter 0.55 mm; pale bluish-white, hemispherical with regular covering of 
spicules. 

Larva. First instar. 0.75 mm in length, 0.25 mm in width; oval in shape; head tan 
brown, lying well beneath prothorax; uniform pale yellow; fringed with fine white setae. 

Second instar. 3 mm by 1.5 mm; similar to first but color straw-yellow; dorsal vessel 
shows as dark green middorsal line; middorsal line white, not extending onto thoracic or 
anal segments; four orange-red spots laterally, two at center of body and two on penul- 
timate abdominal segment. 

Third instar. 5 mm by 2.5 mm; similar to second but color greenish yellow with lateral 
spots brown; middorsal line extends along entire abdomen; setal fringe 0.55 mm in length. 

Fourth instar. 8 mm by 4 mm, greatest width being at metathorax; dark green, paling 
to edges; middorsal line wholly white, or with pattern of brownish purple spots, three 
centrally and one on penultimate abdominal segment. 

Fifth instar. 12 mm by 6 mm; dark green; middorsal line creamy white, along entire 
body length; setal fringe 2 mm in length (Fig. 12). 

Pupa. 9 mm in length, 5 mm in width; similar to P. intensa; smooth, not hirsute; dark 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 4-13. Philiris spp., larvae and pupae: 4 & 5, P. moira; 6 & 7, P. helena; 8 & 
9, P. agatha; 10 & 11, P. intensa; 12 & 13, P. ziska. 


green with middorsal pattern of chocolate-brown flecks on white on abdominal segments 
1-4; same pattern repeated at apex of mesothorax and both areas surrounded by pale 
yellow; spiracles white. Supported by cremaster and fine silk girdle (Fig. 13). Duration, 
12 days. 


BioLocy. The foodplant is Malaisia scandens (Lour.) Bl. (Moraceae) 


VOLUME 38, NUMBER 1 pas 


which grows to about 5 m tall and is common along creeksides or road 
verges around Bulolo (altitude 700 m). Young leaves are soft and pale 
green. Old leaves are dark green and extremely brittle to the touch. 
They are ovate and may reach a length of 150 mm. The tree is a 
sprawling species which sends out long adventitious shoots. The small 
fruits are soft and red and are borne in many clusters along the 
branches. 

Adults of P. ziska are often fairly common where the foodplant 
grows, especially alongside creeks. 

Eggs are laid anywhere on the underside of new or old leaves, some- 
times five on a leaf. The newly emerged larva eats the top of the egg 
and leaves a white ring of shell which remains attached to the leaf. At 
all stages the larvae feed on the lower epidermis of the leaf and leave 
the upper epidermis as windows of tissue. They will not accept any 
other related plant species. The larval ground color is a perfect mimic 
of a vein. They were seen to be attended by small brown ants. 

Pupation is always on the upperside of the leaf along the mid-vein 
just before it joins the petiole. Up to 12 pupae have been found dis- 
persed throughout one branch. The ground color is a perfect match of 
that of the leaf, and the brown markings edged with yellow resemble 
blemishes that are typically found on the leaves of various moraceous 
tree species. 

P. ziska appears to be prone to attack by small, black chalcid wasp 
parasitoids. For example, one larva collected in its fourth instar ceased 
feeding and two 2 mm-long wasp larvae emerged to spin their cocoons 
beneath it before the larva died. Some pupae found had also died from 
what appeared to be a fungal disease. 


DISCUSSION 


A number of adults of each species have been reared and compared 
with material in the British Museum (Natural History) collection. In 
all cases males and females compared favorably with the pairings pre- 
sented in the collection (as figured by D’Abrera, 1977). Representatives 
of each species have been placed in the collection of the Insect Farming 
and Trading Agency in Bulolo. 

Comparison of the early stages, especially the morphology of the 
pupae, shows that P. helena and P. agatha are closely related as are 
P. ziska and P. intensa. The pupa of P. moira resembles more the 
latter pair than those of P. helena or P. agatha but is very distinct in 
that it is prominently hirsute, not smooth as in P. ziska and P. intensa, 
and its maculation is different. It is very similar to that of P. innotata 
and also that of P. kapaura Tite. Pupae of the latter species I have 


22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


found attached to saplings of a species of fig with large (500 mm 
diameter), hirsute leaves. Therefore, P. moira, P. innotata and P. ka- 
paura form a closely related group of species. The similarities between 
the larval morphologies and foodplant relations of P. ziska, P. intensa 
and P. moira (together with P. innotata and P. kapaura), however, 
suggest that these butterflies may also belong to the same species group. 
Females of P. moira are virtually indistinguishable from those of P. 
ziska. 

It is noteworthy that I have found P. moira larvae feeding on Ficus 
semivestita Corner (Moraceae), near Bulolo, as well as Ficus calopilina 
Diels. The latter is the foodplant recorded for the species by Forbes 
(UST) 


ACKNOWLEDGMENTS 


My thanks to to Kenneth Airy Shaw of Kew Herbarium, England for his assistance in 
the identification of the foodplants and to Don Sands of C.S.I.R.O., Queensland, Australia 
for kindly sending me separates of his papers. 


LITERATURE CITED 


AirY SHAW, H. K. 1980. The Euphorbiaceae of New Guinea. Kew Bull. Additional 
series VIII. 243 pp. 

COMMON, I. F. B. & D. F. WATERHOUSE. 1981. Butterflies of Australia. Second revised 
edition, Angus and Robertson, Sydney. 682 pp. 

D’ABRERA, B. 1977. Butterflies of the Australian Region. 2nd edition. Landsdowne, 
Melbourne. 415 pp. 

FORBES, G. R. 1977. The life history and polymorphic female of Philiris moira Grose- 
Smith (Lepidoptera: Lycaenidae) from Papua New Guinea. J. Aust. Entomol. Soc. 
16:273-275. 

GRESSITT, J. L. & N. NADKARNI. 1978. Guide to Mt. Kaindi: background to montane 
New Guinea ecology. Wau Ecology Institute Handbook No. 5. 135 pp. 

SANDS, D. P. A. 1979. New species of Philiris Réber (Lepidoptera: Lycaenidae) from 
Papua New Guinea. J. Aust. Entomol. Soc. 18:127-183. 

SANDS, D. P. A. 1980. The identity of Philiris nitens Grose-Smith (Lepidoptera: Ly- 
caenidae), with description of a new subspecies from Papua New Guinea. Aust. 
Entomol. Mag. 6:81-86. 

SANDS, D. P. A. 1981. New species of Philiris Réber (Lepidoptera: Lycaenidae) from 
mainland New Guinea. J. Aust. Entomol. Soc. 20:89-96. 


Journal of the Lepidopterists’ Society 
88(1), 1984, 23-31 


COURTSHIP BEHAVIOR OF THE GULF FRITILLARY, 
AGRAULIS VANILLAE (NYMPHALIDAE) 


RONALD L. RUTOWSKI AND JOHN SCHAEFER 
Department of Zoology, Arizona State University, Tempe, Arizona 85287 


ABSTRACT. The courtship behavior of the Gulf Fritillary, Agraulis vanillae L., is 
described from motion picture records of successful and unsuccessful courtships between 
free-flying males and tethered virgin females. During most but not all courtships the 
male performs a previously undescribed wing clap display in which he alights next to 
the female and repeatedly claps his wings together, often catching the female’s antenna 
between his wings during each clap. In the discussion it is suggested that this display 
presents chemicals signals to the female and has evolved in response to female choice 
for males that clearly announce their species identity to the female. 


In recent years substantial literature has developed concerning the 
behavior and ecology of the heliconiine butterflies (for review: Brown, 
1981). Surprisingly, only two papers (Crane, 1955; Gilbert, 1976) have 
dealt specifically with the mating behavior of heliconiines in spite of 
a growing amount of information on the nutritious materials passed by 
males to females during copulation and the female’s use of these ma- 
terials in oogenesis (Boggs and Gilbert, 1979; Boggs, 1981). The only 
courtship description per se for a heliconiine is to be found in Crane’s 
paper on Heliconius erato Hewitson. Information on courtship behav- 
ior is essential in any attempt to evaluate the selective consequences 
of inter- and intra-specific variation in male’s abilities to produce ac- 
cessory gland secretions during copulation (Rutowski et al., 1983). 

The following study expands our knowledge of the behavior of hel- 
iconiines by describing the courtship of a member of this subfamily 
that is common in the southern United States, the Gulf Fritillary 
(Agraulis vanillae Linnaeus). Particular attention during this study was 
directed at describing a previously unreported display performed by 
A. vanillae males and its role in successful courtship. The discussion 
will focus on the potential functions of this unique display. 


METHODS 


Observations were made on a population of A. vanillae in suburban 
Tempe, Arizona. Preliminary studies in the spring of 1980 led to in- 
tensive study from March to June in 1981. From about 0800 h to 1600 
h, adults of A. vanillae frequently visit Lantana spp. (adult nectar 
source) and Passiflora spp. (larval foodplant). Virgin females were ob- 
tained from larvae and eggs collected on Passiflora spp. either in the 
field or in cages in the laboratory. Larvae were reared to adulthood in 
translucent plastic shoeboxes (9 x 16 x 30 cm) on cuttings of Passi- 
flora. The shoeboxes were kept near a window which exposed the 


24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


larvae to a normal light-dark regime. Temperature and humidity were 
not controlled. 

To observe courtship, newly emerged, virgin females no more than 
four to five days old (most were one day old) were tethered (for tech- 
nique: Rutowski, 1978) and placed on a conspicuous perch on a Pas- 
siflora plant. Usuaully within a few minutes, passing males approached 
and courted the female. Courtships, both successful and unsuccessful, 
were filmed at 24 and 70 frames per sec with a Beaulieu 4008 ZM 2 
super-8 movie camera. 


RESULTS 
Successful Courtship 


By separating males and females immediately after coupling, twen- 
ty-five successful courtships (leading to copulation) were filmed using 
ten females. No single female was used to film more than three court- 
ships. Of these 25 courtships, one involved two males and so, was 
disregarded, leaving 24 for the detailed analysis that gave rise to the 
description that follows. 

Courtship began when either the male alit next to the female or the 
female began a flutter response as the male approached. Both were 
often preceded by a brief period during which the male hovered about 
15 cm over the female before descending and initiating physical con- 
tact. Once the male alit he positioned himself with his wings open, his 
head close to that of the female, and the long axis of his body forming 
about a 45 degree angle with that of the female (Fig. 1). Once in this 
position the male began what will hereafter be referred to as the wing 
clap display. During this display the male’s body remained in position 
but he repeatedly clapped his wings shut and then quickly reopened 
them. Between claps the wings were opened to about 90 degrees rel- 
ative to one another. It was typical that in the position assumed by the 
male, the female’s antenna on the side next to the male was laid back 
between the male’s wings and was caught between them during each 
clap. During wing clapping and in some instances even before the 
male alit, the male’s claspers were visibly spread. Just as, or before, the 
male ceased clapping his wings he began probing by curling his ab- 
domen toward the female’s hindwings and attempting to insert its tip 
up between her closed wings. Coupling occurred between the female’s 
hindwings where its occurrence could not be closely monitored. How- 
ever, it is assumed that either when it occurred or shortly thereafter 
the male became still briefly before he began slowly waving his wings 
and moving to a position facing away from the female while still 


VOLUME 38, NUMBER 1] 25 


Yeon ~ 





Fic. 1. Above: an A. vanillae male (left) as he appears when courting a female 
(right) with a wing clap display. Below: the same pair except that the male has now 
begun probing. In both figures, note that the female’s antenna on the side next to the 
male is laid back between his wings. 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Temporal structure of successful courtship in Agraulis vanillae. 


Time event occurs before end of courtship 


Courtships with wing clap display Courtships without wing clap display 

Event x + SD (s) range (sec) n x + SD (s) range (sec) n 

First contact by 6 10.9 = 5.1 PPro) UT 8.04 + 3.67 5438-153 6 

Last contact by 6 Chil ae BOX AAVaIS.j3 — IT Oj) as 2/7 1.8-7.46 6 
6 begins wing clap 

display ees) 22 2 1.92-15.8 17 — —- 


Ol 


6 begins probing 3.538 + 3.18 0.5-13.0 14 2.97 + 1.46 1.46-5.53 
6 ends wing clap 

display Wess) ae Pho) OS f—alPAish 7 — = a 
6 stops moving 0 0 


coupled. Wing waving often evolved directly into strong wing flapping 
by the male in an effort to initiate a post-nuptial flight. 

This sequence of events was observed in 14 of the 24 courtships 
filmed. In another three records the male broke contact with the perch 
and alit again at least once but no more than twice. In one case the 
male performed a bout of probing while perched the first time. After 
alighting a final time, all of these males performed wing claps before 
probing. 

A more striking variant of the courtships described above was ob- 
served in seven courtships in which the wing clap display was com- 
pletely omitted. In these cases the male simply began probing imme- 
diately after alighting for the last time as they did in some of the 
courtships with wing claps but in these cases the males were successful. 
Hence, the wing clap display is not a requisite part of successful court- 
ship. 

Females opened and closed or fluttered their wings in only eight of 
the 24 courtships. In six of these the female performed a single flutter 
before or at the time of first contact by the male. If the male made 
repeated contact each contact often elicited a single flutter. Twice 
flutters were observed during the wing clap display and twice after 
the male had become still. Fluttering in immediate response to male 
contact was observed in three of the seven courtships in which males 
did not wing clap. 

The temporal structure of courtships in which the male performed 
a wing clap display is shown in Table 1. In these summaries, there are 
some courtships in which the first contact was also the last; hence, some 
data points were used in the summaries for both events. Also, for 
convenience and summary purposes, we regarded the last time the 
male initiated probing as the time when the male began probing. This 
was not true of three courtships. The sample size of the summary of 


VOLUME 38, NUMBER 1 ; PLT) 


30 


Frequency (°%o) 


O — 
O ICO> 2007300; "400 >400 


Duration (msec) 


Fic. 2. The frequency distribution of duration for 132 wing claps. See text for details. 


this event is less than that for the other events because in three cases 
the angle of filming did not permit viewing of the beginning of prob- 
ing. 

Courtships with the wing clap display have an average duration of 
about 11 sec and wing clapping begins within a second after the male 
alights for the last time. Probing usually began in the second before 
the display ended and slow wing waving followed coupling within 2 
sec. 

Table 1 also summarizes the temporal structure of courtships in 
which the male did not perform a wing clap display. The sample size 
for some events is less than seven, either because the film record began 
after the event occurred or because the filming angle prohibited view- 
ing the event. Statistical comparisons of the time of occurrence of 
events held in common between the courtships with and without the 
wing clap display revealed that none of the differences were signifi- 
cant. In particular, neither the time of first contact nor the time of last 
contact differed from one type of courtship to the next. Factors con- 
tributing to this lack of differences were the high variances associated 
with the times of occurrence and the small sample sizes for courtships 
without the wing clap display. 

The average duration of the wing clap display was 4.73 + 3.34 sec 
(n = 17). Fig. 2 shows the average duration of each clap from the time 
the wings start to close from their spread position until they start to 
close for the next clap. The data are taken from 132 wing claps per- 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


formed in five displays of 6, 12, 16, 41, and 57 claps, respectively, and 
filmed at 70 frames per sec. The histogram shows a mild but distinct 
bimodality. This reflects the fact that during the display the temporal 
patterning of the claps may take one of three forms: (1) a series of 
long claps, (2) a series of short claps, and (3) a series of alternating 
long and short claps. From the films we also measured that, regardless 
of overall clap duration, the wings were closed an average of 26.4 + 
9.37 msec (n = 127). Hence, the difference in duration between long 
and short claps is due primarily to a difference between them in the 
time the wings are open. 

Most courtships were filmed at 24 frames per sec which made count- 
ing of individual wing claps difficult; thus, we were unable to obtain 
a good average for the number of wing claps per courtship. However, 
using a mean wing clap display duration of 4.73 sec and an average 
of 165 msec per clap, it may be estimated that in the typical courtship 
the male performs about 30 wing claps. 


Unsuccessful Courtships 


Courtships that did not lead to copulation were studied in less detail 
than those that lead to copulation. This was because unsuccessful court- 
ships were more variable and difficult to characterize. In addition many 
of the film records of unsuccessful courtship were incomplete. How- 
ever, 20 complete records were obtained and are summarized here. In 
five of these courtships the male contacted the female and hovered 
over her briefly before departing. In the other 15 the male alit next to 
the female at least once during the courtship. Of the males that alit, 
four departed without performing a wing clap display while the other 
11 performed the display at some point in the courtship. Two males 
actually probed before wing clapping while three did not begin prob- 
ing until they had wing clapped for some time. The other six males 
left the female after wing clapping without probing. During these 
courtships the females either did nothing (10 cases), fluttered the wings 
(8 cases), or assumed a posture like the pierid mate refusal posture 
(Obara, 1964) with the wings spread and the abdomen held perpen- 
dicular to the plane of the wings (2 cases). When females fluttered or 
spread their wings it could be seen that the pair of glands associated 
with the tergites at the end of the abdomen were periodically everted. 
Whether or not this occurs when the wings are closed was not deter- 
mined. 

In summary, courtship terminated before coupling either because 
the male left before an attempt (males did not probe in 16 cases) or 
because the female did not behave in a way that permitted coupling 
(4 cases). 


VOLUME 38, NUMBER 1 29 


DISCUSSION 


The wing clap display performed by males of Agraulis vanillae is 
clearly an important part of successful courtship. Its form is reminis- 
cent of displays performed by males of some other nymphalids. The 
male of the grayling (Hipparchia (=~Eumenis) semele Linnaeus) after 
alighting next to a perched female moves so that he is face to face with 
the female and, by rocking forward with the wings open, catches the 
female’s antennae between his forewings. The male then closes his 
wings and rocks gently back, drawing the female’s antennae across 
patches of scent scales on the male’s forewings (Tinbergen et al., 1942). 
During the courtship of the great spangled fritillary (Speyeria cybele 
Fabricius), the male assumes a position perched next to the female like 
that seen in A. vanillae, and “‘at intervals he would suddenly open and 
close his wings’ (Clark, 1932:110). The interval between these open- 
ings and closings becomes less as the courtship progresses. Magnus 
(1950) reports that males of the fritillary (Argynnis paphia Linnaeus) 
when perched alongside and facing a female, clap their wings in such 
a way that the female’s head and antennae are caught between them. 
In Heliconius erato, the closest relative of A. vanillae that has been 
carefully studied, the male persists in flapping his wings after alighting 
and while moving into position for coupling (Crane, 1955). Perhaps 
this behavior was the one from which the wing clapping display of A. 
vanillae was evolutionarily derived. 

Indications are that the wing clapping display is involved in pre- 
senting chemical signals to the female. Obviously, the male’s position 
behind and to the side of the female is poor for presentation of visual 
stimuli, and the fact that the female’s antenna is between the male’s 
forewings during the display suggests that the display is designed to 
deliver a chemical signal. This conclusion is reinforced by the obser- 
vation that androconia that are likely to be scent-producing are found 
on the dorsal surface of the male’s forewings along several of the veins 
(M,, M,, M,, Cu,, Cu,, 2A; Muller, 1877). In addition there are large 
apparently glandular structures associated with the internal faces of 
the claspers that might produce a chemical signal. The claspers are 
often spread or opened during most of the courtship. Experiments are 
planned to determine the roles of these structures in the courtship 
behavior of A. vanillae. 

Rutowski (1983) has recently discussed the selection pressures that 
have led to the evolution of species-specific male courtship displays in 
butterflies. A similar analysis of the courtship of A. vanillae suggests 
that selection for signals that announce the species identity of the male 
to the female has been of particular importance. The nutrient invest- 
ment of A. vanillae males and the general ecology and behavior of 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


this species is not so different from that of the other species of butter- 
flies as to suggest that they led to the evolution of the wing clapping 
display (Rutowski et al., 1983). In addition, although A. vanillae 
females may occasionally approach and chase males as observed in 
other species (Crane, 1955; Rutowski, 1980; Rutowski et al., 1981; Wik- 
lund, 1982), it does not seem that problems of sexual discrimination 
for females are any more severe than they are in Heliconius erato, 
which like A. vanillae, is essentially sexually monomorphic with re- 
spect to visual characters but whose courtship lacks the wing clap 
display (Crane, 1955). 

Species discrimination would seem, however, to pose a potential 
problem for A. vanillae females. A field guide to the butterflies of 
North America (Pyle, 1981) reveals that there are at least eight but- 
terflies sympatric with A. vanillae that are very similar in color and 
markings. These species include Danaus gilippus Cramer, D. plexip- 
pus Linnaeus, Dryas julia Fabricius, Marpesia petreus Cramer, Li- 
menitis archippus Cramer, Dione moneta Hiibner, Anaea floridalis 
Johnson and Comstock, and A. andria Scudder. Given that an A. va- 
nillae female is likely to be approached and courted by males of any 
of these species, selection might favor those females that prefer con- 
specific males that perform a display or offer a signal clearly indicative 
of their species identity. While it is known that the danaids have their 
own unique male courtship behavior (hairpencilling: Brower et al., 
1965; Pliske, 1975) the other species are not behaviorally well-known 
enough to assess the extent to which their courtships are different from 
that of A. vanillae. 

A final question about the wing clap display concerns the fact that 
it is not performed in all successful courtships like the wing spread 
display sometimes performed by males of the pierid, Nathalis iole 
Boisduval (Rutowski, 1981(83)). Why do some females accept a male 
without the performance of the display? Obviously, females vary in 
receptivity, and some will accept males without the display. These may 
be males with such potent chemical signals that they are able to sat- 
isfactorily stimulate the female without the display. In any event it 
seems from both successful and unsuccessful courtships that males will 
attempt copulation without wing clapping, which in turn suggests that 
the display has certain costs for males. Just what these costs are or 
might be is currently not clear (although, see: Burk, 1982). 


ACKNOWLEDGMENTS 
Thanks to Michael Boppré for showing us the location of androconia in A. vanillae, 


to Mark Newton for assistance in filming courtship, and to the National Science Foun- 
dation for financial support (Grant No. BNS 80-14120). 


VOLUME 38, NUMBER 1 ; 3] 


LITERATURE CITED 


Bocecs, C. L. 1981. Selection pressures affecting nutrient investment at mating in 
heliconiine butterflies. Evolution 35:931-940. 

Boccs, C. L. & L. E. GILBERT. 1979. Male contribution to egg production in butterflies: 
evidence for transfer of nutrients at mating. Science 206:83-84. 

BROWER, L. P., J. V. Z. BROWER & F. P. CRANSTON. 1965. Courtship behaviour of the 
queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 50:1-39. 

Brown, K. S., JR. 1981. The biology of Heliconius and related genera. Ann. Rev. 
Entomol. 26:427-—456. ; 

BuRK, T. 1982. Evolutionary significance of predation on sexually signalling males. 
Florida Entomol. 65:90-104. 

CLARK, A. H. 1932. The butterflies of the District of Columbia and vicinity. Smithson- 
ian Institution, U.S. National Museum Bulletin 157. U.S. Government Printing Office, 
Washington. 

CRANE, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara 
Hewitson, with special reference to the social use of color. Zoologica 40:167-196. 

GILBERT, L. E. 1976. Postmating female odor in Heliconius butterflies: a male-con- 
tributed antiaphrodisiac? Science 193:419—420. 

Macnus, D. B. E. 1950. Beobachtungen zur Balz und Eiablage des Kaisermantels 
Argynnis paphia L. (Lep., Nymphalidae). Zeit. Tierpsych. 7:435-449. 

MULLER, F. 1877. The scent-scales of the male of Dione vanillae. Kosmos 2:38-41. 
(Translation in: Longstaff, G. B. 1912. Butterfly Hunting in Many Lands. Longmans, 
Green, and Co., London.) 

OparRA, Y. 1964. Mating behavior of the cabbage white Pieris rapae crucivora. II. The 
‘mate refusal posture’ of the female. Dobut. Zasshi 73:175-178. 

PLISKE, T. 1975. Courtship behavior of the monarch butterfly, Danaus plexippus L. 
Ann. Entomol. Soc. Amer. 69:143-151. 

PYLE, R. M. 1981. The Audubon Society Field Guide to the North American Butterflies. 
A. A. Knopf, Inc., New York. 916 pp. 

RUTOWSKI, R. L. 1978. The form and function of ascending flights in Colias butterflies. 
Behav. Ecol. Sociobiol. 3:163-172. 

RUTOWSKI, R. L. 1980. Courtship solicitation by females of the checkered white but- 
terfly, Pieris protodice. Behav. Ecol. Sociobiol. 7:113-117. 

RUTOWSKI, R. L. 1981(83). Courtship behavior of the dainty sulfur butterfly, Nathalis 
iole, with a description of a new facultative male display. J. Res. Lepid. 20:161- 
169. 

RUTOWSKI, R. L. 1983. The wing waving display of Eurema daira males (Lepidoptera, 
Pieridae): its structure and role in successful courtship. Anim. Behav. 31:985—989. 

RUTOWSKI, R. L., C. E. Lonc, L. D. MARSHALL & R. S. VETTER. 1981. Courtship 
solicitation by Colias females (Lepidoptera: Pieridae). Amer. Midl. Nat. 105:334- 
340. 

RUTOWSKI, R. L., M. NEWTON & J. SCHAEFER. 1988. Interspecific variation in the size 
of the nutrient investment made by male butterflies during copulation. Evolution 
34:708-713. 

TINBERGEN, N., B. J. D. MEEUSE, L. K. BOEREMA & W. W. VAROSSIEAU. 1942. Die 
Balz des Samtfalters, Euwmenis (=Satyrus) semele (L.). Zeits. Tierpsych. 5:182-226. 

WIKLUND, C. 1982. Behavioural shift from courtship solicitation to mate avoidance in 
female ringlet butterflies (Aphantopus hyperanthus) after copulation. Anim. Behav. 
30:790-793. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 32-39 


CHECKLIST OF MANITOBA BUTTERFLIES 
(RHOPALOCERA) 


PAUL KLASSEN 
Box 212, Elm Creek, Manitoba 


ABSTRACT. A list of butterflies (Rhopalocera) occurring in Manitoba is compiled 
from records of resident and non-resident collectors, published literature, museums, uni- 
versity collections and the author’s collection. 


It has been forty years since the last published checklist of Manitoba 
butterflies (Rhopalocera) by G. Shirley Brooks in “A Revised Check 
List of the Butterflies of Manitoba” (1942). Since that list is out-dated 
and not readily available, the present list has been prepared, including 
a number of species not previously recorded. 

Many parts of Manitoba have been collected very sparingly, and I 
am afraid the habitat will be destroyed before these areas have been 
studied. There is very little virgin prairie left in this province, and 
some of that is not accessible to collectors. Some species in this habitat 
are threatened. Most of the province, however, is largely undeveloped, 
and there are large tracts of virgin forests, marshes, bogs, taiga and 
tundra untouched by the bulldozer. This will pigea lly remain so for 
a long time. 

It is hoped that this checklist will encourage more study of the 
fascinating butterfly fauna of Manitoba. Any comments and criticism 
of this list and the notes following it will be appreciated. 

The sequence of taxa follows the order of the Miller and Brown 
Catalogue/Checklist (1981), and the species are numbered accord- 


ingly. 
DISTRIBUTION 


Most of Manitoba is covered by boreal forest including many lakes, 
rivers and bogs. The southern part, especially toward the west, consists 
of grasslands changing to an aspen parkland region farther north. The 
area bordering the coast of Hudson Bay contains some tundra. 

For practical reasons the following definitions are used: 


FN = Far North. An area just southwest of Hudson Bay. Here Churchill and vicinity 
have been collected quite intensively and most far north records are from here. 

N = North. Northern third of the province excepting the far north. This area consists 
of boreal forest with make lakes, rivers and bogs. Not much collecting has been done in 
this area. 

NW = Northwest. The western half of “N”’. 

NE = Northeast. The eastern half of “N”. 

C = Central. The middle third of the province running north and south. Geographi- 
cally this area is like the north. Very little collecting has been done here. 


VOLUME 38, NUMBER 1 a 


WC = West Central. The western half of “C”. 

EC = East Central. The eastern half of “C”’. 

S = South. The southern third of the province. 

SW = Southwest. The western half of “S”. This area consists of dry prairie in the 
southwest turning to moist prairie farther north and east. A large part of this is in the 
parkland or transition zone and is broken up by the Turtle Mountain in the extreme 
south and the Riding and Duck Mountains to the north. Lake Manitoba is east of Riding 
Mountain. Most of this area is agricultural land with very little virgin prairie left. 

SE = Southeast. The eastern half of “S’’. This area consists of boreal forest in the north, 
mixed forest farther south with moist prairie along the Red River valley. The southern 
end of Lake Winnipeg is included in this area. Most of the prairie is now in agriculture. 

G = General Distribution. Covers the whole province. 

Note: Only those areas for which there are actual records of butterflies have been 
listed. Some species probably cover a much larger area than is indicated in the checklist 
below. 


CHECKLIST OF MANITOBA BUTTERFLIES 
(RHOPALOCERA) 


Hesperiidae Latreille 
Epargyreus Hiibner 


clarus clarus (Cramer)—S, C Ta. 
Thorybes Scudder 

pylades (Scudder)—G (except FN) 48. 
Erynnis Schrank 

icelus (Scudder & Burgess)—G (except FN) 83. 

brizo brizo (Boisduval & Leconte)—S, C 84a. 

juvenalis juvenalis (Fabricius)—S 85a. 

martialis (Scudder)—SE 92. 

lucilius (Scudder & Burgess)—S 96. 

persius persius (Scudder)—SW, NE, FN 99a. 
Pyrgus Hiibner 

centaureae freija (Warren)—SE, WC, N, FN 100a. 

communis (Grote)—S 104. 
Pholisora Scudder 

catullus (Fabricius)—S eS: 
Carterocephalus Lederer 

palaemon mandan (Edwards)—S, C 120a. 
Ancyloxypha 

numitor (Fabricius)—S, C 142. 
Oarisma Scudder 

poweshiek (Parker)—S 144. 

garita (Reakirt)—S 145. 
Thymelicus Hiibner 

lineola (Ochsenheimer)—SE 150. 
Hesperia Fabricius 

uncas uncas Edwards—SW 156a. 

comma assiniboia (Lyman)—S 158b. 

c. borealis Lindsey—NE, FN 158d. 

ottoe Edwards—S 160. 

leonardus Harris—SE 161. 

pawnee Dodge—SW 162. 

dacotae (Skinner)—S 169. 

sassacus manitoboides (Fletcher)—SE 171b. 


nevada (Scudder)—SW £78. 


34 


Polites Scudder 

coras (Cramer)—S 

themistocles (Latreille)—S 

mystic dacotah (Edwards)—S, WC 
Atrytone Scudder 

logan lagus (Edwards)—SW 
Poanes Scudder 

hobomok (Harris)—S 
Euphyes Scudder 

ruricola metacomet (Harris)—S, C 
Atrytonopsis Godman 

hianna hianna (Scudder)—S 
Amblyscirtes Scudder 

hegon (Scudder)—SE 

vialis (Edwards)—S 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


174. 

179. 
181b. 
189b. 

1973 
217b. 
219a. 


235. 
245. 


Papilionidae Latreille 


Papilio Linnaeus 
polyxenes asterius Stoll—SE 
bairdii Edwards—SW 
kahli F. & R. Chermock—SW 


machaon hudsonianus Clark—SW, WC, N 


cresphontes Cramer—S 


glaucus canadensis Rothchild & Jordan—G 


troilus troilus Linnaeus—SE(?) 


308a. 
308. 
306. 
310b. 
314. 
320b. 
325a. 


Pieridae Duponchel 


Pieris Schrank 
protodice Boisduval & Leconte—S 


occidentalis occidentalis Reakirt—S, FN 


napi oleracea Harris—G 
rapae (Linnaeus)—S, FN 
Euchloe Hiibner 


ausonides mayi F. & R. Chermock—G 


olympia (Edwards)—SW 

Colias Fabricius 
philodice philodice Godart—S, C 
eurytheme Boisduval—S, FN 


alexandra christina Edwards—SW, WC 


hecla hela Strecker—FN 

boothii Curtis—FN 

nastes moina Strecker—FN 
gigantea gigantea Strecker—FN 
g. mayi F. & R. Chermock—SW 


pelidne pelidne Boisduval & Leconte—FN 


interior interior Scudder—S, WC 
palaeno chippewa Edwards—FN 
cesonia (Stoll) —SW 
Eurema Hiibner 
mexicana (Boisduval)—SW 
Nathalis Boisduval 
iole Boisduval—S 





334. 
335a. 
336d. 

338. 


84le. 
344. 


351a. 

on2. 
355e. 
357b. 

358. 
360c. 
362a. 
362c. 
368a. 
364a. 
365a. 
368a. 


380. 


389. 


Lycaenidae Leach 


Feniseca Grote 


tarquinius tarquinius (Fabricius)—S 


Lycaena Fabricius 
xanthoides dione (Scudder)—S 


39 la. 


395b. 


VOLUME 38, NUMBER 1 


hyllus (Cramer)—S 
epixanthe michiganensis Rawson—SE 
dorcas dorcas Kirby—G 
helloides (Boisduval)—S 
Harkenclenus dos Passos 
titus titus (Fabricius)—S 
Satyrium Scudder 
acadica acadica (Edwards)—SE 
a. watrini (Dufrane)—SW 
edwardsii (Grote & Robinson)—S_ - 
calanus falacer (Godart)—S 
liparops fletcheri (Mitchener & dos Passos)—S 
Incisalia Scudder 
augustus augustus (Kirby)—S, C, N 
polios polios Cook & Watson—S, C, N 
henrici henrici (Grote & Robinson) —SE 
niphon clarki Freeman—S, C 
eryphon eryphon (Boisduval)—N 
Strymon Hiibner 
melinus humuli (Harris)—S 
Everes Hiibner 
comyntas comyntas (Godart)—S, C 
amyntula aibrighti Clench—SW, WC, N, FN 
Celastrina Tutt 
ladon lucia (Kirby)—G (except FN) 
l. argentata (Fletcher)—SW 
Glaucopsyche Scudder 
lygdamus couperi Grote—G (except FN) 
l. afra (Edwards)—-SW 
Plebejus Kluk 


Argyrognomon scudderii (Edwards)—SW, WC, FN 


a. nabokovi Masters—SE 

melissa melissa (Edwards)—-SW 

m. samuelis Nabokov—SE 

saepiolus amica (Edwards)—G 
optilete yukona (Holland)—C, N, FN 
franklinii franklinii (Curtis)—FN 

f. lacustris (Freeman)—C, N 

f. rustica (Edwards)—S 


Heliconiidae Swainson 


Agraulis Boisduval & Leconte 
vanillae incarnata (Riley)—SW 


Nymphalidae Swainson 


Euptoieta Doubleday 
claudia (Cramer)—S 
Speyeria Scudder 


cybele pseudocarpenteri (F. & R. Chermock)—S 


aphrodite aphrodite (Fabricius)—SE 

a. manitoba (F. & R. Chermock)—S 
idalia (Drury)—S 

edwardsii (Reakirt)—SW 

callippe calgariana (McDunnough)—S 
atlantis atlantis (Edwards)—SE 

a. hollandi (F. & R. Chermock)—S, WC 


565d. 
566a. 
566d. 

567. 

569. 
o72p. 
o7 4a. 
o74c. 


36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a. dennisi dos Passos & Grey—SW 574u. 
mormonia eurynome (Edwards)—SW 576i. 
Boloria Moore 
eunomia dawsoni (Barnes & McDunnough)—G 578c. 
selene atrocostalis (Huard)—S, WC, FN 579f. 
bellona bellona (Fabricius)—S, WC, N 580a. 
frigga saga (Staudinger)—-G 58 la. 
improba improba (Butler)—FN 582a. 
polaris stellata Masters—FN 585b. 
freija freija (Thunberg)—G 586a. 
titania boisduvalii (Duponchel)—FN 589a. 
t. grandis (Barnes & McDunnough)—G (except FN) 589c. 
chariclea arctica (Zetterstedt)—N 590a. 
Chlosyne Butler 
gorgone carlota (Reakirt)—S 605b. 
nycteis nycteis (Doubleday)—S 606a. 
n. reversa (F. & R. Chermock)—SW 606c. 
harrisii harrisii (Scudder)—S 607a. 
h. hanhami (Fletcher)—S, WC 607c. 
Phyciodes Hiibner 
tharos tharos (Drury)—G 623b. 
batesii (Reakirt)—S, C 624. 
Euphydryas Scudder 
phaeton phaeton (Drury)—SE 635a. 
Polygonia Hiibner 
interrogationis (Fabricius)—S 636. 
comma (Harris)—S 637. 
satyrus neomarsayas dos Passos—S 638b. 
faunus faunus (Edwards)—S, WC 639a. 
gracilis (Grote & Robinson)—N, FN 643. 
progne (Cramer)—S, N, FN 645. 
Nymphalis Kluk 
vau-album j-album (Boisduval & Leconte)—S 646a. 
californica californica (Boisduval)—S 647a. 
antiopa antiopa (Linnaeus)—G 648a. 
milberti milberti (Godart)—S, WC, FN 649b. 
Vanessa Fabricius 
virginiensis (Drury)—S, FN 650. 
cardui (Linnaeus)—S, WC, FN Gal. 
atalanta rubria (Fruhstorfer)—S, C, N 653a. 
Junonia Hiibner 
coenia Hiibner—S 656. 
Limenitis Fabricius 
arthemis arthemis (Drury)—SE 663a. 
s. rubrofasciata (Barnes & McDunnough)—S, C 663b. 
archippus archippus (Cramer)—S, WC 664a. 


Satyridae Boisduval 
Lethe Hiibner 


anthedon Clark—S TU 

eurydice eurydice (Johansson)—S 718a. 
Euptychia Hiibner 

cymela cymela (Cramer)—S, C 723a. 
Coenonympha Hiibner 

inornata inornata Edwards—SW, C 728d. 


i. benjamini McDunnough—S 728e. 


VOLUME 38, NUMBER 1] OU 


Cercyonis Scudder 


pegala olympus (Edwards)—S 732e. 
Erebia Dalman 
rossi ornata Leussler—FN 737a. 
disa mancinus Doubleday & Hewitson—G 738a. 
discoidalis discoidalis (Kirby)—G 7Ala. 
theano sofia Strecker—FN 742a. 
epipsodea freemani Ehrlich—SW, WC 744b. 
Neominois Scudder 
ridingsii ridingsii (Edwards)—SW . 748a. 
Oeneis Hiibner 
macounii (Edwards)—S (ile 
chryxus calais (Scudder)—C 752b. 
uhleri varuna (Edwards)—-SW 753a. 
alberta alberta Elwes—S 754a. 
bore ssp.—FN 756. 
jutta ascerta Masters & Sorensen—SE 757b. 
j. ridingiana F. & R. Chermock—SW, WC Tne: 
j. harperi Chermock—N, FN Told. 
melissa semplei Holland—FN 758c. 
polixenes polixenes (Fabricius)—FN 759a. 


Danaidae Duponchel 


Danaus Kluk 
plexippus (Linnaeus)—S, WC 760. 


NOTES 


C. palaemon mandan—Type-locality—“Lake Winnipeg’, restricted to Pine Ridge by 
F. M. Brown and L. Miller, is common in most wooded areas of southern Manitoba. 

T. lineola, first recorded from Manitoba in the early 1970’s, is now firmly established 
in Winnipeg and east of there (Preston and Westwood, 1981). 

H. comma borealis from Churchill, should perhaps have another subspecific name. 

P. asterius polyxenes is rare in southeastern Manitoba. 

P. kahli—Type-locality—“Riding Mtns., Man.”, is found mostly in the Riding Moun- 
tain and Duck Mountain area, but some are found as far east as the Red River. There 
seems to be some intergradation between this and the latter species and P. machaon. 

E. ausonides mayi—Type-locality—“Riding Mtns., Manitoba”. 

C. hecla hela—Type-locality—“‘above Fort Churchill”. 

C. nastes moina—Type-locality— ‘above Fort Churchill”. 

C. g. giganiea—Type-locality—‘“west coast of Hudson Bay above Fort York”’. 

C. g. mayi—Type-locality— “Riding Mtns., Manitoba”’. 

L. d. dorcas—Type-locality— ‘Lat. 54°”, restricted to The Pas, Manitoba, by Ferris. 

H. t. titus—In southwestern Manitoba some specimens perhaps belong to immaculo- 
sus. 

S. a. acadica flies in the southeast and watrini in the southwest. 

S. liparops fletcheri—Type-locality—““Manitoba’”’. 

C. ladon argentata—Type-locality—‘“Cartwright, Manitoba’, flies in southwestern 
Manitoba, while lucia is found in most of the rest of the province. 

G. lygdamus afra—Type-locality— “Deer River country ’’, restricted to vic. Brandon, 
Man. by F. M. Brown, flies in southwestern Manitoba with couperi in the rest of the 
province. They are quite hard to tell apart as they are variable in size and in the ventral 
spots and color. 

P. argyrognomon scudderii—Type-locality—“Lake Winnipeg, Manitoba’, flies in 
western Manitoba. The bands of submarginal orange lunules, both ventral and dorsal, 
are on the average more complete than in nabokovi which is found in the southeastern 
part of the province. The subspecies are very variable and so difficult to tell apart. 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


P. m. melissa flies in south and central Manitoba with samuelis in the southeastern 
corner. 

P. f. franklinii is found in the Churchill area. 

P. f. lacustris—Type-locality—“Norway House’, is in central Manitoba. 

P. f. rustica occurs in southern Manitoba. 

S. aphrodite manitoba—Type-locality—“Sand Ridge’, which is east of Riding Moun- 
tain. These formerly went under the name of mayae and occupy most of southern 
Manitoba. S. a. aphrodite is in the extreme southeast. 

S. a. atlantis is sometimes found in the extreme southeast. 

S. a. hollandi—Type-locality— ‘Riding Mtns., Manitoba’, flies in most of southern 
Manitoba. 

S. a. dennisi—Type-locality—““Beulah, Manitoba’, closely resembles lais from Sas- 
katchewan. It was known by that name for some time. It probably intergrades with that 
subspecies. In the Riding Mountain area, hollandi flies in the wetter areas and dennisi 
in the drier, more open areas, but adults feed at flowers in the same places. Should these 
belong to different species? 

B. polaris stellata—Type-locality—“Churchill, Manitoba’, flies in the Churchill area 
in odd-numbered years. 

B. titania grandis flies in southern and central Manitoba. 

B. t. boisduvalii from the north is abundant at Churchill. 

C. nycteis reversa—Type-locality—“Riding Mountains, Manitoba’, refers to most 
specimens from Manitoba, however some are like subspecies nycteis in facies. 

C. harrisii hanhami—Type-locality— ‘Bird Hill, near Winnipeg, Manitoba’, flies in 
southern Manitoba. There seems to be some intergradation with the subspecies harrisii, 
as some specimens are like the ones from Ontario in facies. 

L. arthemis rubrofasciata—Type-locality— “Manitoba, Saskatchewan, Alberta’, is 
common in southern Manitoba with some arthemis found in the eastern part, where 
they intergrade. 

L. eurydice, formerly known as transmontana in this area, is quite distinct as the 
ground color very pale, almost white in some specimens, as compared to the dark spec- 
imens found in eastern Ontario. 

C. i. inornata—Type-locality—“Lake Winnipeg’, emended to “Saskatchewan River 
between Lake Winnipeg and The Pas, Man.”, by F. M. Brown, flies mostly in the 
parklands area and benjamini on the prairies in southern Manitoba. C. ochracea probably 
does not fly in the province. 

E. rossii ornata—Type-locality—“Churchill”, is abundant in the Churchill area most 
years. 

E. theano sofia—Type-locality—“Fort Churchill, Manitoba” was formerly known as 
canadensis, is locally common at Churchill most years. 

O. bore ssp., flies at Churchill in even-numbered years. It is quite rare most years, but 
locally more common, in some. It is quite variable and has a darker ground color than 
bore hanburyi from Baker Lake, N. W. Territories, Canada. 

O. jutta ascerta flies in eastern Manitoba. It is dark and the orange bands are less 
developed or even lacking in some males. It is found in the odd-numbered years with 
the rare exception. 

O. j. ridingiana—Type-locality—“Riding Mountains, Manitoba’’, is found mostly in 
even-numbered years in western Manitoba, but some fly every year. The orange bands 
are well developed. 

O. j. harperi—Type-locality—“Gillam, Manitoba’, is a little smaller than the two 
preceding subspecies. It is quite variable with the orange bands in some females well 
developed to faint in others. It resembles alaskensis. It is common at Churchill every 
year. 

Some butterflies have, over a number of years, been taken very rarely in Manitoba. 
The following are probably strays from the south: P. cresphontes, P. t. troilus, C. cesonia, 
E. mexicana, N. iole, A. vanillae, S. idalia and N. californica. 

There are also some species that, although rare, apparently breed in the province. 
Some of these may be seen to be more common after the areas have been more exten- 


VOLUME 38, NUMBER 1 39 


sively collected. Here is a list of these: P. catullus, O. poweshiek, H. ottoe, H. dacotae, 
H. nevada, A. logan, P. bairdii, E. olympia, L. epixanthe, I. eryphon, S. melinus, S. 
edwardsii, S. callippe, E. phaeton, P. gracilis, J]. coenia and N. ridinsii. 

A small number have been included that maybe should be deleted from the list. Papilio 
bairdii is included based on records from Beulah and Birtle and records of bairdii ore- 
gonia from Beulah. The author suspects that these may be misidentified specimens of 
machaon or kahli. The latter is quite variable. C. boothii, C. pelidne and B. improba 
have been recorded from “north Manitoba”. C. boothii and B. improba could occur 
northwest of Churchill and C. pelidne could be found east of there. B. chariclea is 
recorded from Kettle Rapids. Formerly Boloria titania from Manitoba were called chari- 
clea titania. As there is no proven reason to the contrary, the above specimens are all 
included in the checklist. 

The following species, included in older lists, have been deleted: H. comma manitoba, 
no records for Manitoba. 

P. zelicaon probably does not occur in the province. The records possibly refer to 
machaon or forms of kahli. 

E. ausonides coloradensis is supposed to fly in southeastern Manitoba. I cannot see 
any difference between the mayi, type-locality, “Riding Mtns., Man.’’, and the ausonides 
from the rest of the province. 

P. zephyrus recorded from Aweme and Beulah probably were misidentified Polygonia. 

S. cypris = ethene and S. a. columbia included in older lists probably are S. a. mani- 
toba, which they closely resemble. 

S. lais, included in old lists flies in Saskatchewan and Alberta and intergrades with 
dennisi in Manitoba. 

S. calanus calanus recorded as calanus is deleted as the subspecies that flies in the 
province is falacer. 

S. heathii, also omited, because it is an aberration of the latter. 

S. liparops strigosa does not occur in the province. Although some specimens of fletch- 
eri from Manitoba closely resemble strigosa with no orange spots on the fore-wings, these 
occur in the same populations together with specimens having orange patches covering 
one-half of the front wings. This subspecies is very variable. 

Mitoura spinetorum probably does not fly in Manitoba. 


ACKNOWLEDGMENTS 


Many thanks go out to all the following, who sent in data and helped in other ways: 
George T. Austin, Patrick J. Conway, Richard E. Gray, W. W. Gregory, R. J. Heron, 
Ronald R. Hooper, Brian McKillop and William B. Preston of the Manitoba Museum of 
Man and Nature, David Parshall, James D. Reist, Oakley Shields and Jim Troubridge, and 
to the personnel of the University of Manitoba and the Canada Agriculture Research 
Station. 


LITERATURE CITED 


BROOKS, G. SHIRLEY. 1942. A check list of the butterflies of Manitoba. Can. Entomol. 
74:31-36. 

MILLER, LEE D. & F. MARTIN BROWN. 1981. A Catalogue/Checklist of the Butterflies 
of America North of Mexico. The Lepid. Soc. Memoir No. 2. 280 pp. 

PRESTON, W. B. & A. R. WESTWOOD. 1981. The European Skipper, Thymelicus lineola 
(Lepidoptera: Hesperiidae), in Manitoba and Northwestern Ontario. Can. Entomol. 
113:1123-1124. 

WALLIS, J. B. 1927. A Colour Key to the Manitoban Butterflies. Nat. Hist. Soc. Man. 
31 pp. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 40-46 


THE LIFE HISTORY AND BEHAVIOR OF 
EPIMARTYRIA PARDELLA (MICROPTERIGIDAE) 


PAUL M. TUSKES 
1444 Henry St., Berkeley, California 94709! 


AND 


NORMAN J. SMITH 
2192 Jenni Ave., Sanger, California 93657 


ABSTRACT. Adults of Epimartyria pardella (Walsm.) are rather sessile and exhibit 
a clumped distributional pattern. Moths are active during the day and usually closely 
associated with liverworts. Larvae from eggs deposited in the lab feed on liverworts. 
There are three larval instars and in captivity 1.75 years were spent in the larval stage. 
Collection of wild larvae suggest that 2 years are also required to complete development 
under natural conditions. 


The family Micropterigidae is recognized as the most primitive group 
of Lepidoptera known. The adult moths are the only Lepidoptera with 
functional mandibles which they use for feeding on pollen. Microp- 
terigidae are aglossate, jugate moths whose closest relatives are believed 
to be the Heterobathmiidae. Chapman (1917) and Hinton (1946) placed 
the Micropterigidae in their own order, the Zeugloptera, because of 
the primitive characters the larvae express, but Common (1970), Kris- 
tensen (1971) and Richards and Davies (1959) treated the Zeugloptera 
as a suborder of Lepidoptera. Fossil micropterigids in lower Cretaceous 
amber indicate that relatively little change has occurred in the group 
during the last 185 million years (Whalley, 1977, 1978). 

In the United States this unique suborder is represented by the new 
world genus Epimartyria Walsm. (1898) that consists of two species. 
A great deal of work has been done on the systematics and evolutionary 
status of the micropterygids (Hinton, 1958; Common, 1975; Heath, 
1976; Whalley, 1978; Kristensen and Nielsen, 1979), but observations 
dealing with their behavior and habitat are for the most part lacking. 
In this paper, information is presented on the biology and habitat of 
Epimartyria pardella (Walsm.). 

The type series of E. pardella consists of five specimens which were 
collected near the coast in southern Oregon during early June 1872. 
The description that Walsingham (1880) published is brief and accom- 
panied by a color illustration. The moth (Fig. 1) has a wingspan of 10 
to 11 mm. The forewing is metallic brown with three distinctive gold 
spots, while the hind wings are only metallic brown. The fringe of both 


1 Present address: 7900 Cambridge 141G, Houston, Texas 77054. 


VOLUME 38, NUMBER 1 Al 


fore and hindwings is yellow and brown. The abdomen and thorax are 
gray-brown; the legs and a portion of the head are golden yellow. From 
the head to the posterior tip of the abdomen the moth measures just 
under 3.5 mm. 


Last Instar Larva 


Head. Length 0.5 mm, diameter 0.27 mm. Brown. Antennae prominent, trisegmented 
and situated on small tubercles located on dorsal lateral portion of head (Fig. 3). Stem- 
mata with 5 facets and located at the base of the antenna. Labrum simple with a pair 
of trisegmented palpi. Mandibles simple and dark brown. 

Body. Length 4.3 to 4.6 mm; width 1.4 mm; height 1.2 mm. The body tapering at 
both ends with highest and widest point at abdominal segment 4. Dorsal and lateral 
surface brown to dark brown, ventral surface light brown. Prothoracic shield with 10 
peg-like setae, 8 on the anterior and lateral border and 2 dorsally. Prothorax distinctly 
narrower than mesothorax. Mesothorax with 8 setae, 6 on dorsal and lateral anterior 
portion of gray brown pigmented area, and 2 just ventral to this pigmented area. Setae 
of metathoracic segment similar to those of mesothorax except subdorsal seta is greatly 
reduced in size. All thoracic segments have additional small micro-seta just dorsal to each 
true leg. True legs brown, with 3 segments and simple claw. Abdominal segments (A) 
Al to A8 (and T2 and T3) with sawtooth-shaped knobs which form a dorsal and lateral 
ridge, areas between ridges concave. The middorsal area concave and small dark depres- 
sion occuring on posterior of segments T2 to A8. Segments Al to A8 each with one dorsal 
seta (0.18 mm) atop dorsal ridge. Segments Al to A8 with reduced, almost microscopic 
subdorsal seta (0.04 mm) and prominent lateral seta (0.12 mm) on lateral ridge. Dorsal, 
subdorsal and lateral setae occur in brown pigmented area which has rough and wrinkled 
appearance. Dorsal and lateral intersegmental area constricted and may contain series 
of 8 to 20 microscopic dots. Ventral to lateral ridge, cuticle smooth and light brown. 
Series of brown dots form pattern around fixture that usually support a small seta. Conical 
ventral “prolegs” occur on segments Al to A8 and small sclerotized protuberance appears 
on ventral surface of each. Segments A9 and A10 fused and with enlarged simple sucker. 
Spiracles posterior and ventral to lateral setae. Head diameter of first and second instar 
larvae 0.11 and 0.22 mm, respectively. 


Habitat 


Observations were made in Prairie Creek State Redwood Park, 
Humboldt County, California. All locations where adults were ob- 
served or captured were within a few km of the ocean and at relatively 
low elevations. Although some moths were found along creeks and 
moist hillsides in the redwood-fir forest, the preferred habitat appears 
to be steep-walled, moist canyons near the coast which are dominated 
by ferns and bryophytes (Fig. 6). The prominent bryophytes that are 
associated with the adults and larvae are, Conecephalum conium, Pel- 
lia sp., Hookeria lucens, and Atrichum undulatum. Other plants in 
the immediate area include: bracken fern, Pteridium aquilinum; sword 
fern, Polystichum minitum; deer fern, Blechnum spicans; and five- 
finger fern, Adiantum pedatum. 

Climatic conditions in this area are moderate and stable. Weather 
records from Prairie Creek Campground, located about 4 km east of 
the beach at an elevation of 160 m indicated the mean daily temper- 


42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-5. 1, Adult male E. pardella (14x); 2, ova on underside of liverwort thallus 
(8x); 3, last instar larva (9x); 4, cocoon (5x); 5, pupa (9x). 


ature for January (9.5°C) and July (15.5°C) of 1981 differed by only 
6°C. Although the summer months (June to September) are relatively 
dry, approximately 140 cm of rain falls between October and May. 
During the 1980-81 rainy season (October to May) there were 12 days 
when the temperature dropped below O0°C (32°F); the lowest temper- 
ature recorded during that time was —1.5°C (28°F). Barbour et al. 
(1973) suggested that seasonal temperature fluctuation reaches a min- 


VOLUME 38, NUMBER 1 43 





Fic. 6. Habitat of E. pardella in Northern California. 


imum in this area because of off shore upwelling. They indicated that 
the mean monthly air temperature normally changes only a few de- 
grees between the coldest and warmest months, and the ocean tem- 
perature changes very little. 


Adult Behavior 


The flight season begins in late May and continues to early or mid- 
July, with the peak adult density in June. The moths are active during 
the day, generally between 0900 and 1930 h, but this is influenced by 
temperature, humidity, and light intensity. When abundant, adults 
may be observed perched on vegetation; at low densities the best means 
of locating a colony is by sweeping suitable habitat with an aerial net. 
Behavioral notes were made on the activities (in situ) of individual 
moths that were observed from one to seven hours. 

In areas protected from wind, adults frequently perched on the 
upper surface of fern fronds or other plants near patches of liverworts 
growing on canyon walls or beside creeks. Adults exhibit a clumped 
distributional pattern and, where common, densities reached 6 moths/ 
m?. When windy, or if the humidity is low, adults find shelter among 
the moist bryophytes with which they are always closely associated. 
The antennae are held at a 45 to 60 degree angle above the midline 
of the body while the moths are perched (Fig. 1). As they walk, the 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


antennae wave up and down often touching the substrate. The hind 
legs of pardella are almost equal in length to that of the entire moth, 
and are occasionally used to jump or hop a few centimeters. Adults 
may remain motionless for hours and then walk or fly a few centi- 
meters and perch again. During a single two and one half hour obser- 
vation, a moth traveled 25 cm in a sporadic pattern and came to rest 
for the evening less than 5 cm from where it was first observed. Another 
moth less than 50 cm away walked less than 15 cm during this time. 
Most moths in 1981 were observed from one to three hours and trav- 
eled less than 30 cm. In 1982, five moths were carefully observed for 
a total of 29.2 hours. Again, the adults were extremely sessile, often 
remaining for hours in the same position. During the 29.2 hours, 16 
flights were observed with an average distance of 21 cm per flight; 
they walked an average of 17 cm. Moths changed positions to perch 
in sunny locations, to avoid predators, and in the case of females, to 
oviposit. Adults are the prey of various small predators. One moth was 
captured in a spider web and another chased by a small hunting spider 
of the genus Theridion. A third moth was stalked but not captured by 
a small Olympic salamander (Rhyacotriton olympicus). Moths fly when 
disturbed but normally flight is infrequent and brief; the flight pattern 
is fluttery and weak but usually direct. 

Adult micropterigids of other genera are reported to feed on pollen 
rather than nectar and have unique mouthparts. The mandibles are 
well developed, and the hypopharynx is concave on the upper surface. 
As pollen grains are ingested they are ground by the action of the 
mandibles against the hypopharyngeal spines and then digested (Till- 
yard, 1923; Hannermann, 1956). European species have been collected 
at the blooms of many plant species, including: Compositae, Acer, 
Carex, Scrophulariaceae, Quercus, and Ranunculus (Heath, 1960). Al- 
though various Ranunculus, Compositae, and Scrophulariaceae were 
near by and in bloom, no moths were observed at the flowers. Adults 
were frequently observed drinking water. Since they lack a proboscis 
they lower their head to the droplet of water by extending their meso- 
thoracic legs to the side of their body. This lowers the head and raises 
the abdomen, allowing the moth to drink. If deprived of moisture 
moths die in less than two days, but when provided with water, they 
survived in captivity from nine to 18 days, and females deposited ova. 

The only mating pair of moths was found just prior to 1000 h. In 
captivity females laid an average of 8.2 eggs per day. Ova were de- 
posited on the underside of the liverwort thalli singly or in small clus- 
ters containing up to five ova (Fig. 2). The females generally remained 
on the upper surface and would simply swing their abdomen under 
the edge of the thallus to oviposit. 


VOLUME 38, NUMBER 1 45 


Immature Stages 


The ova are flattened, circular and smooth when first deposited but 
become spherical in a short time and are covered with a series of small 
white projections (Fig. 2). The ova are white and measure 0.40 x 0.44 
mm. At 22°C the eggs hatch in 21 days. The first instar larvae emerge 
from the side of the egg and are about 0.75 mm long. They vary from 
light brown to light gray and appear to have the same setal pattern 
and shape as mature larvae but have the ability to flatten themselves 
when at rest. 

Larvae were reared in either a terrarium or petri dishes. Although 
both species of liverwort (Conocephalum and Pellia) were available, 
the larvae showed a marked preference for Pellia, the smaller of the 
two species. Mature larvae are active primarily at night but early instar 
larvae may be active at any time. While feeding, the margin of the 
liverwort is not damaged, rather the underside of the living thallus is 
eaten away but not through. Many micropterigid species feed on bryo- 
phytes, but the work of Luff (1964) and Lorenz (1961) indicates that 
some species do not. 

In captivity the larvae are rather inactive, avoid intense light, and 
are usually found on the underside of the thalli during the day. In the 
field larvae were also found under living thalli during the day. Their 
coloration and size allowed them to blend well with the dead thalli 
which occur under the living growth. As the larva walks the true legs 
grasp the substrate; from above it appears to glide across the surface 
as the rhythmic undulations of the ventral surface are not apparent. 
When disturbed or inactive the head may be withdrawn so that only 
the prothoracic shield is visible; when extended the antennae which 
are located above the eyes are prominent (Fig. 3). 

Unlike the European species which have a one year life cycle (Heath, 
1976; Lorenz, 1961), pardella appears to have a two year cycle. In 
captivity eggs deposited in June 1981 became adults in June 1983. In 
the field, second instar larvae were commonly collected each year 
during the adult flight period. These larvae must represent the off- 
spring from ova deposited the previous year, as reared larvae one year 
old were also in the second instar. Davis (pers. comm.) observed that 
E. auricrinella (Walsm.) from the eastern United States also has a two 
year life cycle. 

Pupation occurs close to the ground among vegetation. The brown 
cocoon, which measures 5.5 x 4.5 mm, is oval, thin walled and tightly 
woven (Fig. 4). Strands of coarse silk attach the cocoon to vegetation. 
The exarate pupa is white to light brown (Fig. 5). 

Based on the illustration of Micropterix calthella (L.) larvae by Lo- 
renz (1961), the larvae of E. pardella exhibit a number of differences. 


46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The setae of calthella are club-shaped and apparently uniform in length. 
The larvae of pardella have peg-shaped setae which vary in length 
according to their location. In addition, the distribution of the larval 
setae and pupal setal patterns also differ. A preserved pupa, cocoon, 
and larvae were deposited in the collection of the California Academy 
of Sciences, San Francisco. 


ACKNOWLEDGMENT 


The research conducted at Prairie Creek Redwood State Park was done under a permit 
from the California Department of Parks and Recreation. We wish to thank the park 
service for their cooperation. We also wish to thank Ann McGowan-Tuskes for two years 
of field assistance and for reviewing the manuscript. 


LITERATURE CITED 


BaRrBour, M. G., R. B. Craic, F. R. DRUSDALE & M. T. GHISELIN. 1973. Coastal 
Ecology: Bodega Head. Univ. of Calif. Press, Berkeley. 

CHAPMAN, T. A. 1917. Micropteryx entitled to ordinal rank; Order Zeugloptera. Trans. 
Entomol. Soc. London 1916:310-314. 

Common, I. F. B. 1970. Lepidoptera. In insects of Australia. Melbourne. Melbourne 
Univ. Press. Pp. 765-866. 

1975. Evolution and classification of the Lepidoptera. — Rev. Entomol. 20: 
183-203. ‘ 

HANNERMANN, H. J. 1956. Die Kopfmuskulatur von Micropteryx calthella L. Mor- 
phologie und funktion. Zool. Jahrb. Anat. 75:177-206. 

HEATH, J. 1960. The foodplants of adult micropterygids. Entomol. Mon. Mag. 95:188. 

1962. The eggs of Micropteryx. Ibid. 97:179-180. 

1976. The moths and butterflies of Great Britain and Ireland. Vol. 1. Pp. 151- 











155. 

HINTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of lepidoptera. Trans. Roy. Entomol. Soc. 
London 97:1-87. 

1958. The phylogeny of the oaneneatd orders. Ann. Rev. Entomol. 3:181—206. 

KRISTENSEN, N. P. 1971. The systematic position of the Zeugloptera in the light of 
recent anatomical investigations. Proc. XIII Int. Cong. Entomol. 1:261. 

KRISTENSEN, N. P. & E. S. NIELSEN. 1979. A new subfamily of micropterigid moths 
from South America. A contribution to the morphology and phylogeny of the Mi- 
cropterigidae, with a generic catalogue of the family (Lepidoptera: Zeugloptera). 
Steenstrupia 5(7):69-147. 

LORENZ, R. E. 1961. Biologie und morphologie von Micropterix calthella (L.). Dt. Ent. 
Z. (N.F.) 8:1-28. 

Lurr, M. L. 1964. Larvae of Micropteryx [sic] (Lepidoptera; Micropterygidae). Proc. 
R. Entomol. Soc. Lond. (C) 29:6. 

RICHARDS, O. W. & R. G. DaAvigs. 1957. In a general textbook of entomology. A. D. 
Imms. London, Methuen. 9th ed. 886 pp. 

TILYARD, R. J. 1923. On the mouth parts of the Micropterygoidea (Lepidoptera). Trans. 
Roy. Entomol. Soc. London 181-206. 

WHALLEY, P. E. S. 1977. Lower Cretaceous Lepidoptera. Nature 266:526. 

1978. New taxa of fossil and recent Micropterygidae with a discussion of their 
evolution and a comment on the evolution of Lepidoptera. Ann. Transvaal Mus. 31: 
71-86. 

WALSINGHAM, T. 1880. On some new and little known species of Tineidae. Proc. Zool. 
Soc. London 83-84. 

1898. Description of a new micropterygid genus and species and a new erio- 

craniad species from N. America. Entomol. Rec. J. Var. 10:161-163. 











Journal of the Lepidopterists’ Society 
88(1), 1984, 47-50 


A NEW ACANTHOPTEROCTETES FROM THE 
NORTHWESTERN UNITED STATES 
(ACANTHOPTEROCTETIDAE) 


DONALD R. DAVIS 


Department of Entomology, Smithsonian Institution, 
Washington, D.C. 20560 


ABSTRACT. Acanthopteroctetes aurulenta Davis, new species, is described from 
Oregon and Utah. Both male and female are illustrated. 


Recent collecting in central Utah by Ronald W. Hodges resulted in 
the discovery of the male of an undescribed species of Acanthopter- 
octetes previously mentioned in the literature (Davis, 1978:96, 129) 
but not named. The availability of both sexes of this species now en- 
ables me to name this insect, which constitutes only the fourth species 
described for the family. 


Acanthopteroctetes aurulenta, new species 


Length of forewings. 6, 7.4 mm; 8, 5.1 mm (Fig. 1). 
Head. Vestiture rough, pale yellowish brown to nearly white. Antennae with 43 seg- 
ments; vestiture of scape extremely rough with prominent pecten of more than dozen 





Fic. 1. Acanthopteroctetes aurulenta, new species. Holotype 6, wing expanse 15 
mm. 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 2-5. Acanthopteroctetes aurulenta, new species, male genitalia: 2, lateral view, 


(J = juxta; Tr = transtilla); 3, ventral view; 4, valva, mesal view; 5, aedoeagus. Scale = 
0.5 mm. 


long whitish hairs extending over eye; flagellum smooth, uniformly banded with white 
and pale brown scales. Haustellum naked except for scattered, fine setae. Maxillary palpi 
greatly lengthened, 5-segmented, geniculate; vestiture white. Labial palpi considerably 
shorter than maxillary palpi, covered with whitish scales. 

Thorax. Pronotum covered with smooth, golden brown scales; central tuft of approx- 
imately one dozen elongate golden hairs present. Forewings uniformly pale golden brown, 


VOLUME 388, NUMBER 1 49 








Fics. 6-8. Acanthopteroctetes aurulenta, new species, female genitalia: 6, ventral 
view (CO = common oviduct, U = utriculus, V = vesicle); 7, dorsal view; 8, vestibulum 
and bursa copulatrix, dorsal view. Scale = 0.5 mm. 


slightly lustrous; R, slightly variable, either connate with R,,;+M, or shortly stalked. 
Hindwings more thinly scaled, uniformly pale gray. Venter of thorax white. Legs mostly 
white; epiphysis absent. 

Abdomen. Sparsely covered with pale golden brown scales above, more whitish be- 
neath. External glands absent. Caudal margin of eighth segment in female with encir- 
cling ring of elongate sensory setae; median setae longest with setae decreasing in length 
ventrally. 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Male genitalia (Figs. 2-5). Uncus slender, minutely bifid, with 5 minute, subapical 
serrations along ventral margin. Ninth segment relatively long cylinder, about twice 
length of uncus, without lateral separation between tegumen and vinculum. Both anterior 
and posterior margins of vinculum deeply excavated. Median process of transtilla with 
3 pairs of ventral serrations. Juxta elongate, length over 2.5 its width; basal half darkly 
sclerotized. Valvae slender, greatest width (at base) 0.25 its length; saccate membrane 
arising from elongate pouch along distal half of cucullus. Aedoeagus elongate, exceeding 
genital capsule in length; prominent cluster of approximately 6 elongate cornuti present. 

Female genitalia (Figs. 6-8). Apex of ovipositor broad, depressed, triangular in outline, 
with approximately 15-17 serrations bordering lateral margins. Posterior apophyses stou- 
ter than anterior pair. Vestibulum enlarged, extremely irregular in outline, and with 
highly folded, thickened walls. Spermatheca with minute spherical vesicle at posterior 
end of elongate, slightly inflated utriculus; spermathecal papillae not sclerotized. Corpus - 
bursae reduced in size, membranous. 

Types. Holotype 6: Head Ephraim Canyon, 10,000-10,300 ft [3049-3140 m], Senpete 
Co., UTAH, 1 Aug 1981, R. W. Hodges, blacklight, USNM 100671. Paratype: Baker, 
Oregon, Spring Creek, 12, 8 Jul 1966. J. H. Baker (USNM). 

Distribution. Northwestern Oregon and central Utah. 


Remarks 


The uniformly light golden brown forewings of A. aurulenta easily 
distinguishes it from the other darker, banded-wing species in the fam- 
ily. This characteristic color pattern has suggested the specific name, 
derived from the Latin aurulentus (golden, ornamented with gold). 
The valvae of A. aurulenta are also unusual in possessing a very dis- 
tinct, thinly sclerotized pocket from which arises the peculiar saccate 
membrane found in all members of the genus. 

The Spring Creek, Oregon habitat can be characterized as a pine- 
sagebrush association with Ceanothus (the host of A. unifascia Davis 
(Davis and Frack, in press)) occurring nearby. The type locality in the 
Wasatch Mountains of central Utah, which has been heavily grazed in 
recent times (D. C. Ferguson, pers. comm.), is an open, subalpine 
plateau. 


ACKNOWLEDGMENTS 


I wish to thank my assistant, Ms. Biruta Akerbergs Hansen, for preparing the illustra- 
tions for this paper, and Dr. Ronald Hodges of the Systematic Entomology Laboratory, 
USDA, for his efforts in collecting this species. 


LITERATURE CITED 


Davis, D. R. 1978. A revision of the North American Moths of the superfamily Erio- 
cranioidea with the proposal of a new family, Acanthopteroctetidae (Lepidoptera). 
Smithsonian Contr. Zool., No. 251, 131 pages, 344 figs. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 51-56 


TWO INTERESTING ARTIFICIAL HYBRID CROSSES 
IN THE GENERA HEMILEUCA AND ANISOTA 
(SATURNIIDAE) 


RICHARD STEVEN PEIGLER! 
303 Shannon Drive, Greenville, South Carolina 29615 


AND 


BENJAMIN D. WILLIAMS 
The Lawrence Academy, Groton, Massachusetts 01450 


ABSTRACT. Two crosses were reared to the adult stage with saturniid moths from 
different areas of the United States. These were Hemileuca lucina 6 x H. nevadensis 2 
reared in Massachusetts and Texas on Salix, and Anisota senatoria 6 x A. oslari 2 reared 
in Connecticut on Quercus coccinea. Larvae and adults of both crosses were interme- 
diate. Descriptions and figures of the hybrids are given. Several isolating mechanisms 
between the parent species were tested and are discussed. 


Dozens of artificial crosses in the Saturniidae have been successfully 
reared since the previous century, but virtually all of these have in- 
volved species of the subfamily Saturniinae. This paper deals with two 
remarkable crosses obtained by the junior author utilizing small satur- 
niid moths belonging to the subfamilies Hemileucinae and Ceratocam- 
pinae.? In both crosses, species native to the Southwest were reared in 
the Northeast and females from those rearings attracted congeneric 
diurnal males native to the Northeast. The species involved were 
Hemileuca lucina Henry Edwards, H. nevadensis Stretch, Anisota 
senatoria (J. E. Smith) and A. oslari W. Rothschild. For information 
on the adult morphology, wing pattern, immature stages, hostplants, 
reproductive behavior, and geographical distributions of these four 
parent species, the reader is referred to works by Ferguson (1971) and 
Riotte and Peigler (1981). 


Hemileuca lucina 6 x H. nevadensis 2 


In mid-September 1977 two virgin females of H. nevadensis (stock 
from Escondido, San Diego Co., California) were placed on twigs of 
Salix gracilis Anderess at the edge of a wet meadow in Groton, Mid- 
dlesex Co., Massachusetts, which supports a sizable population of H. 
lucina. The emergence time of the reared H. nevadensis in Groton 
coincides with the flight time of H. lucina, i.e., mid-September through 
early October. The females emitted pheromone, and males of H. lu- 
cina were attracted. We assumed that pheromone from wild females 


‘Museum Associate in Entomology, Los Angeles County Museum of Natural History. 


52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of H. lucina was also present in the air, and therefore we concluded 
that males of H. lucina may not discriminate between pheromones of 
the two species. No behavioral isolation was observed as is often the 
case in achieving cross-matings of Saturniidae; there was no hesitation 
by the males nor resistance to them by the females. 

Each female produced an egg ring around a twig, each ring con- 
sisting of ca. 100 eggs. The egg is the overwintering stage in most 
species of this genus. One egg ring was sent to the senior author. The 
following spring eclosion of both egg rings was near 100 percent, and 
both authors reared broods successfully to the adult stage. In Groton 
the larvae were reared on Salix gracilis under a cloth bag, surviving 
an unseasonal 25 cm snowfall on 10 May while in the second instar. 
In Brazos County, Texas, the senior author reared his brood on Salix 
sp. (probably nigra Marshall) under cloth bags. Adult emergence in 
1978 in Texas and Massachusetts differed, probably as a result of dif- 
ferences in photoperiod between the two regions where the pupae were 
kept. In Texas males emerged 21 July through 24 August peaking in 
the middle of August; females appeared during the second half of 
August and early September. The hybrid brood in Massachusetts yield- 
ed males from 6 September to 8 October and females mostly during 
the second week of October. A brood of pure H. nevadensis (Escon- 
dido, California) reared alongside the hybrids in Texas produced adults 
of both sexes in September, too late to permit attempts to backcross 
the hybrids, but coinciding with the emergence pattern of pure H. 
nevadensis in Massachusetts as mentioned above. 

Hybrid females of both broods were sterile, based on the fact that 
their abdomens appeared to contain few or no ova. In both broods 
most adults expanded their wings normally after emerging, but some 
specimens, especially among females, failed to spread their wings par- 
tially or totally. This problem is encountered in several species of the 
genus with reared material and is not considered to indicate reduced 
viability resulting from hybridization. 

Only the final instar larva is described, this one showing greatest 
divergence from parent species, but H. nevadensis and H. lucina hard- 
ly differ structurally. Pupal differences were difficult to find between 
the parent species also, so that those given below may not be reliable. 
Adults of this group do not exhibit sexual dimorphism; thus, the sexes 
are not described separately. 


Description 
Mature larva. Integument intermediate: black with numerous dull white oval flecks,* 


many converging but all individually distinct except around spiracles; in H. lucina flecks 
smaller and more widely separated; in H. nevadensis flecks larger and converging to 


VOLUME 38, NUMBER 1 53 





Fics. 1-8. 1 & 2, hybrid pair of Hemileuca lucina 6 x H. nevadensis 2; 3 & 4, pair 
of Anisota senatoria from Pomfret, Connecticut; 5 & 6, hybrid pair of Anisota senatoria 
6x A. oslari 2; 7 & 8, pair of Anisota oslari from Santa Cruz Co., Arizona. 


o4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


form yellowish white areas on integument, especially dorsally. Two dorsal rows of tufted- 
spine scoli stramineous with black tips as in both parents. Subdorsal and subspiracular 
branched scoli black with whitish tips in hybrid and both species. 

Pupa. Anterior rim of each abdominal segment wider as apparently for H. nevadensis. 
Cremaster with stouter curved spines as in H. lucina; H. nevadensis apparently with 
thinner, straighter spines, and possibly fewer than in H. lucina. Head and thoracic 
characters of both parent species and hybrid indistinguishable. 

Adult (Figs. 1 & 2). Elongated white scales on meso- and metathorax sparsely distrib- 
uted among black scales, these white scales more numerous than in H. lucina but much 
less numerous than in H. nevadensis. Whitish band of forewing agreeing with that of 
H. lucina by being wider on ventral side than on dorsal side, but more like H. nevadensis 
by having crenulate outer margin curving parallel with outer wing margin. Discal mark 
in hindwing containing white slit as in father species, this mark often solid black in H. 
nevadensis. Black portions of wings more opaque and coal-black than either parent 
species. 


Anisota senatoria 8 x A. oslari 2 


A freshly emerged female of A. oslari was permitted to emit pher- 
omone in an exposed location at Pomfret, Windham County, Con- 
necticut, during a clear, cool, and windy day in mid-July 1978. The 
undersized moth had been reared on scarlet oak (Quercus coccinea 
Muenchh.) from eggs received from Madera Canyon, Arizona the pre- 
vious year. Anisota senatoria flies in southern New England from mid- 
June through mid-July, whereas reared specimens of A. oslari have 
emerged from mid-July through mid-August. Males of A. senatoria 
seek females from ca. 1130 to 1530 h EST, and the circadian flight 
time of A. oslari is also known to be during midday hours. A male of 
A. senatoria arrived but had considerable difficulty locating the female 
due to gusty wind. He persisted for ca. 1 h before making physical 
contact, at which time copulation readily occurred. Attempts to obtain 
this cross the previous year had apparently failed due to the normally 
larger size of the females of A. oslari, which prevented the males of 
A. senatoria from achieving copulation. 

After mating, the female of A. oslari oviposited freely. Eclosion of 
the eggs was virtually 100 percent. The hybrid larvae were vigorous, 
and several were reared to maturity under cloth bags on scarlet oak. 
The following year the female hybrids emerged during the last few 
days of May, whereas their male siblings appeared from 9 June through 
19 July. Females were apparently sterile, having shrunken abdomens 
as mentioned under the previous cross. 


Description 


Mature larva. Head brown with bold black markings on each side (head of A. senatoria 
solid black; head of A. oslari solid brown; the two-colored head of hybrids remarkable 
because all known species of Anisota have solid colored heads in all instars). Prothoracic 
tergite black as in A. senatoria. Body color black with bold orange stripes on sides and 
two broken orange stripes on top. Anal plate and anal prolegs orange with black markings 
(solid black in A. senatoria, solid brown in A. oslari). Pattern of spines on body more as 


VOLUME 38, NUMBER Il 55 


in A. senatoria but size and arrangement of spines on anal plate intermediate between 
parent species. Median caudal spine long as in A. oslari. 

Male (Fig. 5). Overall appearance strikingly intermediate. Wingshape as in A. sena- 
toria but large size as in A. oslari. Ground color dark purplish brown, forewings having 
brownish orange overtones. Postmedian line weak; barely discernable transparent patch 
in forewing (absent in A. oslari, well-developed in A. senatoria). White discal mark 
large. Forewing with sparse sprinkling of dark spots. Outer margins of hindwings straight. 

Female (Fig. 6). Intermediate in most characters. Wingshape closer to A. oslari. Ground 
color light brownish orange with pinkish suffusion in postmedian area as in father species 
and on hindwing as in mother species. Postmedian line weak in forewing, very faint in 
hindwing. White discal mark surrounded by purple as in A. senatoria. Forewing with a 
few dark spots. 


DISCUSSION 


Hybridization experiments such as these provide data on isolating 
mechanisms and degree of phylogenetic divergence. Aside from the 
obvious one of allopatry, other isolating mechanisms tested by these 
crosses include mechanical, behavioral, viability of immature stages, 
and fertility of adult hybrids. Remarks on each of these were given 
above for both crosses. The differing emergence times between the 
sexes of an individual hybrid brood were proposed by Peigler (1981) 
as an isolating mechanism, because this reduces frequency of F, or 
backcross matings when hybrid broods are produced in nature (when 
primary isolating mechanisms fail). This phenomenon, now widely rec- 
ognized in hybrid Lepidoptera, is well illustrated in the two present 
crosses. We use the term “isolating mechanism”’ in the traditional sense 
as did Solignac (1981), notwithstanding the valid arguments put forth 
by Key (1981) that several independent principles are encompassed by 
the term. 

Genetic compatibility between two taxa, which is to some extent 
correlated with phylogenetic divergence, falls along a continuum. The 
pairs of species in the present study are demonstrated to have an in- 
termediate affinity when compared to the following two extremes. 
Minimal compatibility of parent species would be seen if eggs fail to 
eclose or larvae die in the first instar. This was demonstrated by the 
cross Hemileuca nuttalli (Strecker) ¢ x H. eglanterina (Boisduval) 
in the studies of Collins and Tuskes (1979), which might be expected 
because the parent species are sympatric. On the other hand, what 
appears to be total genetic compatibility in Saturniidae is illustrated 
by crosses (both reciprocals) between the Indian Antheraea roylei Moore 
and the Chinese A. pernyi (Guérin-Méneville). The parent species have 
chromosome numbers of n = 30 and n = 49 respectively, and the hy- 
brid (n = 30) has been reared through more than 20 generations, main- 
taining its increased vigor over the parent species (Jolly, 1974, 1981). 
Most known crosses of Lepidoptera result in more or less vigorous F, 
hybrid adults with reduced fertility, especially in females. 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


It is our hope that this paper will encourage lepidopterists to exploit 
every opportunity to achieve interspecific matings of species that they 
rear. When fertile eggs and viable larvae result, records and descrip- 
tions should be kept, results published, and material deposited in mu- 
seums. 


ACKNOWLEDGMENTS 


We are grateful to J. Steve McElfresh of San Diego, California, for supplying eggs of 
H. nevadensis and A. oslari. Our figures were made by Thomas Marion Hill of Green- 
ville, South Carolina. Drs. W. D. Winter, Jr. and Joseph E. Eger, Jr. made color photo- 
graphs of living larvae of the hybrids and/or pure species which aided formulation of 
the larval descriptions. Material supplied by Earll M. Brown of San Diego also was useful 
in this study. Specimens of both crosses, including the four hybrids figured, have been 
deposited in the Los Angeles County Museum of Natural History, and a pair of the 
Hemileuca cross is in the American Museum of Natural History. 


LITERATURE CITED 


CoLLins, M. M. & P. M. TuskEs. 1979. Reproductive isolation in sympatric species of 
dayflying moths (Hemileuca: Saturniidae). Evolution 33:728-7383. 

FERGUSON, D. C. 1971. Bombycoidea, Saturniidae (in part), in R. B. Dominick et al., 
The moths of America north of Mexico, fasc. 20:2A:153 pp., 11 col. pls., E. W. 
Classey, London. 

Jotty, M. S. 1974. Discovery of new field of tasar on oak and its impact on national 
economy. Central Tasar Res. Sta., Ranchi, Bihar, India. 4 pp. 

1981. Distribution and differentiation in Antheraea species (Saturniidae: Lep- 
idoptera), pp. 1-14 in S. Sakate & H. Yamada eds., Study and utilization of non- 
mulberry silkworms. Symposium in 16th Internat. Congr. Entomol., August 1980, 
Kyoto, Japan. (12) + 78 pp. 

Key, K. H. L. 1981. Species, parapatry, and the morabine grasshoppers. Syst. Zool. 30: 
425-458. 

LEMAIRE, C. 1978. Les Attacidae americains ... The Attacidae of America (=Satur- 
niidae), Attacinae. C. Lemaire, Neuilly. 238 pp., 49 pls. 

PEIGLER, R. S. 1981. Demonstration of reproductive isolating mechanisms in Callosa- 
mia (Saturniidae) by artificial hybridization. J. Res. Lepid. 19:72-81. 

RIOTTE, J. C. E. & R. S. PEIGLER. 1981. A revision of the American genus Anisota 
(Saturniidae). J. Res. Lepid. 19:101-180. 

SOLIGNAC, M. 1981. Isolating mechanisms and modalities of speciation in the Jaera 
albifrons species complex (Crustacea, Isopoda). Syst. Zool. 30:387—405. 

TUSKES, P. M. 1976. A key to the last instar larvae of West Coast Saturniidae. J. Lepid. 
Soc. 30:272-276. 





* Lemaire (1978:23) explained in detail why the name Ceratocampinae is to be used instead of Citheroniinae. 
* Tuskes (1976) stated that these flecks are circular, but in all material we have seen, consisting of several species of 
the genus, these are distinctly oval. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 57-59 


SPERMATOPHORE PERSISTENCE AND MATING 
DETERMINATION IN THE GYPSY MOTH 
(LYMANTRIIDAE)! 


CYNTHIA R. LOERCH AND E. ALAN CAMERON 


Department of Entomology, The Pennsylvania State University, 
University Park, Pennsylvania 16802 


ABSTRACT. Spermatophores were detectable in all female gypsy moths dissected 
within 1.5 h following inception of copulation. After 1.5 h, the percentage of detectable 
spermatophores decreased with time; by 4.5 h, no spermatophore could be detected in 
any mated female moth. The percentages of detectable spermatophores did not differ 
significantly among three gypsy moth populations (laboratory-reared, high and moderate 
density natural populations) for intervals timed from inception of copulation. Examina- 
tion of the bursa copulatrix for the presence of a spermatophore can be useful for rapid 
determination of female gypsy moth mating success. 


The spermatophore of the gypsy moth, Lymantria dispar (L.), is 
formed within the female bursa copulatrix during the first 10 min of 
copulation (Klatt, 1920; Leonard, 1981). It consists of an oval sperm 
sac with a tapered neck that extends into the ductus bursae and a 
proteinaceous mass secreted by the male accessory glands. Proteolytic 
enzymes produced by the female begin to dissolve the spermatophore 
shortly after its formation (Chapman, 1971; Engelmann, 1970). 

However, little is known of the fate of the gypsy moth spermato- 
phore between formation and disintegration. Taylor (1967) reported 
that the spermatophore disintegrates within one or two hours of cop- 
ulation but did not state whether this is time accrued from inception 
or termination of copulation. The distinction is essential since copula- 
tion averages 60-73 min (range = 20-198 min) (Forbush and Fernald, 
1896; Doane, 1968; Waldvogel et al., 1981). Because the gypsy moth 
spermatophore is not persistent, determination of female mating suc- 
cess relies on examining eggs for embryonation several weeks after 
deposition or examining the female reproductive system for the pres- 
ence of sperm (Stark et al., 1974). This paper presents, for the first 
time, data on the persistence of the gypsy moth spermatophore, with 
implications for rapid determination of female mating success. 


MATERIALS AND METHODS 


Laboratory-reared virgin gypsy moths were mated, uninterrupted, 
in arenas described by Waldvogel et al. (1981). The time in copula 
was recorded for each pair. To obtain data on the persistence of the 


‘ Authorized for publication as Paper Number 6368 in The Journal Series of The Pennsylvania Agricultural Experi- 
ment Station. This work was conducted under Experiment Station Project No. 2044, and supported in part by Regional 
Research Project NE-84 (Revised). 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Percentages of spermatophores detectable at intervals timed from inception 
of copulation for three gypsy moth populations: laboratory-reared, and high and mod- 
erate density natural populations. 


Hours 


following % spermatophores detectable 

inception of 

copulation Laboratory-reared High density Moderate density Total 
Le 100.0 (11) 100.0 (9) 100.0 (10) 100.0 (30) 
2.0 81.8 (11) 66.6 (9) 70.0 (10) 73.8 (80) 
ao 70.8 (24) 50.0 (10) 40.0 (10) 59.1 (44) 
3.0 16.7 (24) 44.4 (9) 30.0 (10) 25.6 (48) 
3.5 0.0 (19) - 1 (9) 10.0 (10) 5.3 (38) 
4.0 =) 0 (9) 10.0 (10) 5.3 (19) 
4.5 — (0) — (0) 0.0 (9) 0.0 (9) 


* Values in parentheses are numbers of mated female moths dissected. Percentages did not differ significantly (Chi- 
square test, Fisher’s exact test; P > 0.05) among populations at each time interval. 


spermatophore, females were dissected under a microscope at 30x 
magnification, at intervals timed from inception of copulation. A me- 
dial incision through the abdominal terga provided access to the bursa 
copulatrix. The bursa copulatrix was then dissected in situ and its 
contents compared with those of an unmated female. All matings and 
dissections were performed at room temperature.. These procedures 
were repeated with virgin moths that emerged from pupae collected 
from moderate density (ca. 83000 egg masses/ha) and high density (ca. 
70,000 egg masses/ha) natural populations in Clearfield County, Penn- 
sylvania. Egg mass densities were estimated by the method of Wilson 
and Fontaine (1978). 


RESULTS AND DISCUSSION 


Duration of copulation averaged 87 + 2.3 min for all mated pairs 
(n = 213, range = 22-218 min). The percentages of spermatophores 
that remained detectable at intervals timed from inception of copu- 
lation are presented in Table 1. For each time interval, the percentages 
of detectable spermatophores did not differ significantly among pop- 
ulations (Chi-square test, Fisher’s exact test; P > 0.05). Within 1.5 h 
following inception of copulation, 100% of the spermatophores in all 
populations could be detected. During this period, the shiny white 
spermatophore was visible through the wall of the bursa copulatrix. 
After 1.5 h, the percentage of detectable spermatophores decreased 
with time; the spermatophore was rarely visible through the bursa 
copulatrix wall, and dissection was necessary to determine its presence. 
At 3.5 h following inception of copulation, the spermatophore was 
detectable in less than 12% of the moths examined from any popula- 
tion. By 4.5 h, the contents of the bursa copulatrix of all mated females 
were indistinguishable from those of an unmated moth. 


VOLUME 38, NUMBER 1 59 


These data eliminate the ambiguity arising from Taylor’s (1967) 
report. His observations, if timed from termination of copulation, 
roughly agree with our findings. In other species of Lepidoptera, where 
the spermatophore may persist for several days or more, the bursa 
copulatrix can be examined for the presence of a spermatophore to 
determine whether a female has mated (Burns, 1968; Snow and Car- 
lysle, 1967; Taylor, 1967). Although the gypsy moth spermatophore is 
not persistent, it can be useful for rapid determination of female mat- 
ing success, which may be required in some precopulatory behavioral 
studies. Examination of the bursa copulatrix for a spermatophore is 
highly reliable within 1.5 h following inception of copulation. The 
presence of a spermatophore indicates female mating success and es- 
tablishes that mating occurred less than 4.5 h prior to examination. 
Unfortunately, the absence of a spermatophore does not establish that 
the female gypsy moth is unmated. When no spermatophore is de- 
tectable, the most immediate recourse is examination of the sperma- 
theca for the presence of sperm (Stark et al., 1974). 


ACKNOWLEDGMENTS 


We thank W. Metterhouse and R. Chianese of the New Jersey Department of Agri- 
culture, Division of Plant Industry, for providing laboratory-reared pupae, and S. J. 
Brumbaugh for assisting with mating observation. We also wish to thank P. H. Adler 
and R. O. Mumma, Department of Entomology, The Pennsylvania State University, 
for their helpful criticisms of the manuscript. 


LITERATURE CITED 


BuRNS, J. M. 1968. Mating frequency in natural populations of skippers and butterflies 
as determined by spermatophore counts. Proc. Nat. Acad. Sci. U.S.A. 61:852-859. 

CHAPMAN, R. F. 1982. The insects: structure and function. 3rd Ed. American Elsevier 
Publishing Co., Inc., New York. 992 pp. 

DOANE, C. C. 1968. Aspects of mating behavior of the gypsy moth. Ann. Entomol. Soc. 
Am. 61:768-773. 

ENGELMANN, F. 1970. The physiology of insect reproduction. Pergamon Press Inc., 
New York. 807 pp. 

ForsusH, E. H. & C. H. FERNALD. 1896. The gypsy moth, Porthetria dispar (Linn.). 
Wright and Potter Printing Co., Boston. 495 pp. 

KuLaTT, B. 1920. Beitrage zur Sexualphysiologie des Schwammspinners. Biol. Zentralbl. 
40:539-558. 

LEONARD, D. E. 1981. Bioecology of the gypsy moth, in The gypsy moth: research 
toward integrated pest management, Doane, C. C. & M. L. McManus, eds., U.S. 
Dep. Agric., Tech. Bull. 1584. pp. 9-29. 

SNow, J. W. & T. C. CARLYSLE. 1967. A characteristic indicating the mating status of 
male fall armyworm moths. Ann. Entomol. Soc. Am. 60:1071-1074. 

STARK, R. S., E. A. CAMERON & J. V. RICHERSON. 1974. Determination of mating 
and fertility of female gypsy moths. J. Econ. Entomol. 67:296-297. 

TAYLOR, O. R., JR. 1967. Relationship of multiple mating to fertility in Atteva punc- 
tella (Lepidoptera: Yponomeutidae). Ann. Entomol. Soc. Am. 60:583-590. 

WALDVOGEL, M. G., C. H. COLLISON & E. A. CAMERON. 1981. Durations of pre- 
copulatory periods of laboratory-reared irradiated and non-irradiated male gypsy 
moths. Environ. Entomol. 10:388-389. 

WILSON, R. W., JR. & G. A. FONTAINE. 1978. Gypsy moth egg-mass sampling with 
fixed- and variable-radius plots. U.S. Dep. Agric., Agric. Handbk. 523. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 60-61 


GENERAL NOTES 


INSECT PARASITES AND PREDATORS OF HACKBERRY BUTTERFLIES 
(NYMPHALIDAE: ASTEROCAMPA) 


During the course of collecting and rearing immature stages of hackberry butterflies 
(Nymphalidae: Asterocampa) over the past five years, a number of arthropod parasites 
and predators were encountered. These arthropods have been preserved or their behay- 
iors recorded in hopes of understanding some of the selective pressures which might 
affect the courses of evolution for Asterocampa species. This note is a report of insect 
species which have a greater or lesser effect on survival of the various stages of the 
butterflies. 

Identifications were made by the author with the aid of the cited references and the 
reference collection at Texas A&M University. Help in the collection or identification of 
specimens, or review of the manuscript was provided by L. G. Friedlander, P. Davis, D. 
and D. Paschley, and Drs. H. R. Burke, J. C. Schaffner, and R. Wharton. 

The most frequently encountered parasites of hackberry butterflies are the scelionid 
egg parasites, which occur in all Asterocampa observed. Stink bugs, such as the one 
figured by Langlois and Langlois (1964, Ohio J. Sci. 64:1-11, fig. 11), are the most 
common predators. Only one other insect (at the generic level) has been positively re- 
ported to attack Asterocampa, the larval parasite, Hyposoter fugitivus (Say) (Hym.: 
Ichneumonidae) (Townes, 1945, Mem. Amer. Entomol. Soc. No. 11, Pt. I, pp. 479-925). 


Parasites Reared from Eggs 


1. Hym.: Eulophidae: Tetrastichus spp. (Boucek, 1977, Bull. Entomol. Res. 67:17-80): 
A. clyton (Boisduval & Leconte) egg masses (TEXAS: Brazos Co., 14-VII-79; Menard 
Co., 20-VI-79). 

2. Hym.: Scelionidae: Telenomus spp. (Masner, 1976, Mem. Entomol. Soc. Canada 
No. 97, 87 pp.): A. argus (Bates) egg mass (MEXICO: Oaxaca, 11-VII-81); A. celtis 
(Boisduval & Leconte) eggs (TEXAS: Hidalgo Co., 4-VI-81); A. clyton egg masses (AR- 
IZONA: Pima Co., 23-VIII-80; TEXAS: Brazos Co., 14-VII-79; Menard Co., 20-VI-79; 
San Patricio Co., 3-VI-81; Travis Co., 14-X-77; Waller Co., 8-VII-79; VIRGINIA: West- 
moreland Co., 22-VI-80); A. leilia (Edwards) eggs (TEXAS: Starr Co., 6-VI-81). 


Parasites Reared from Larvae 


1. Dip.: Tachinidae: Euphorocera prob. floridensis Townsend (Aldrich and Webber, 
1924, Proc. U.S. Natl. Mus. 63:1-90; Cole, 1969, The flies of western North America, 
Univ. Calif. Press, Berkeley and Los Angeles, 693 pp.): A. celtis last stage larva (TEXAS: 
Austin Co., 6-VIII-79). 

2. Dip.: Tachinidae: Lespesia prob. aletiae (Riley) (Beneway, 1963, Univ. Kansas Sci. 
Bull. 44:627-686; Cole, 1969, loc. cit.): A. clyton late stage larvae (TEXAS: Gonzales Co., 
30-IX-79). 

3. Hym.: Braconidae: Cotesia spp. (Mason, 1981, Mem. Entomol. Soc. Canada No. 
115, 147 pp.): A. clyton third stage larvae (TEXAS: Gonzales Co., 21-IX-79; Hidalgo 
Co., 138-XI-77; Jeff Davis Co., 15-VIII-81; Uvalde Co., 23-IX-79). 

4. Hym.: Braconidae: Meteorus spp. (Tobias, 1966, Entomol. Rev. 45:348-358): A. 
clyton larvae! (TEXAS: Goliad Co., 6-VI-81; Travis Co., 29-V-78, 20-VII-79). 

5. Hym.: Eulophidae: Elachertus sp. (Peck et al., 1964, Mem. Entomol. Soc. Canada 
No. 34, 120 pp.): A. celtis last stage larva (TEXAS: Travis Co., 21-VI-78); A. clyton 
middle stage larvae (TEXAS: Brazos Co., 14-VII-79; Travis Co., 28-X-77). 

6. Hym.: Ichneumonidae: Microcharops tibialis (Cresson) (Townes, 1969, Mem. Amer. 
Entomol. Inst. No. 18, 307 pp.; Townes and Townes, 1966, Mem. Amer. Entomol. Inst. 
No. 8, 367 pp.): A. clyton third stage larva (LOUISIANA: St. Tammany Parish, 30-III- 
82). 


VOLUME 38, NUMBER | 61 


Parasites Reared from Pupae 


1. Hym.; Chalcidiae: Brachymeria sp. (Howard, 1885, U.S. Dept. Agric., Bur. Ento- 
mol., Bull. No. 5, 47 pp.): A. clyton pupa (TEXAS: Gonzales Co., 15-X-77). 

2. Hym.: Ichneumonidae: Itoplectis conquisitor (Say): A. clyton pupa (TEXAS: Dim- 
mit Co., 21-IV-79). 


Predators 


1. Hem.: Pentatomidae: Apateticus cynicus Say (Slater and Baranowski, 1978, How 
to know the true bugs (Hemiptera-Héteroptera), Wm. C. Brown Co. Publ., Dubuque, 
Iowa, 256 pp.): A. clyton early stage larvae (TEXAS: Travis Co., 26-III, 24-V, 18-X, 31- 
X-77). 

2. Hem.: Pentatomidae: Apateticus lineolatus (Herrick-Schaeffer) (det. J. Eger): A. 
clyton larvae (TEXAS: Cameron Co., 13-III-79). 

3. Hem.: Pentatomidae: Podisus maculiventris (Say) (Slater and Baranowski, 1978, 
loc. cit.): A. clyton early stage larvae (TEXAS: Travis Co., 24-V, 1-VI, 23-28-X-77, 23- 
V-78). 

4. Hem.: Reduviidae: Sinea prob. sanguisuga Stal: A. clyton second stage larva (TEX- 
AS: Travis Co., 29-V-78). 

5. Hem.: Reduviidae: Sinea spinipes (Herrick-Schaeffer) (Slater and Baranowski, 1978, 
loc. cit.): A. clyton early stage larvae (TEXAS: Travis Co., 28-X-77). 

6. Hym.: Vespidae: Polistes exclamans Viereck: A. celtis fifth stage larva (TEXAS: 
Travis Co., 24-IV-78); A. clyton third stage larvae (TEXAS: Travis Co., 25-X-77). 

7. Hym.: Vespidae: Vespula sp.: A. clyton third stage larvae (TEXAS: Travis Co., 31- 
X-77). 


TIMOTHY P. FRIEDLANDER, Department of Entomology, Texas A&M University, 
College Station, Texas 77843. 


‘None were reared from larvae. One female Meteorus was observed to oviposit in A. clyton larvae. One female 
hyperparasite of Meteorus was observed to oviposit in larvae of the same species. One of these hyperparasites was 
reared from Meteorus cocoons taken in close association with A. clyton larvae. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 61-63 


ITHOMIINE BUTTERFLIES ASSOCIATED WITH NON-ANTBIRD 
DROPPINGS IN COSTA RICAN TROPICAL RAIN FOREST 


Adult females of Mechanitis and the allied genus Melinaea (Brown, 1977, Syst. Ento- 
mol. 2:161-197) feed on the fresh droppings of birds (primarily antbirds) that follow 
swarms of army ants through tropical rain forest in Costa Rica (e.g., Ray and Andrews, 
1980, Science 210:1147-1148). These authors conclude that bird droppings resulting from 
birds following army ant swarms provide a predictable nutrient resource for these female 
butterflies, and that the exploitation of this resource may be related, in some yet to be 
studied way, to egg production. In this note I extend the findings of Ray and Andrews 
(op. cit.) to the association of female ithomiines of various genera to fresh droppings of 
bird species not associated with army ant swarms in Costa Rican tropical rain forest. I 
conclude that fresh bird droppings of any kind in such a habitat provide a resource 
exploited by ithomiines on an opportunistic basis. 

Between 1972 and 1980, I conducted several studies of various butterfly species in a 
small parcel of relatively undisturbed mixed primary and secondary-growth tropical rain 
forest (premontane tropical wet forest) at “Finca La Tigra’, near La Virgen (220 m 
elev.), Heredia Province, Costa Rica. The site is about 20 km from the “Finca La Selva” 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Female Godyris zavelata caesiopicta feeding at fresh bird dropping at a light 
gap in the forest habitat at Finca La Tigra in northeastern Costa Rica. 


study site of Ray and Andrews (op. cit.). During this lengthy period, I observed ithomiine 
butterflies feeding on fresh bird droppings splashed on leaves of understory plants, par- 
ticularly along foot paths and light gaps in the forest. This was not a deliberate search 
for butterflies, but rather accidental encounters within an approximately 500-square 
meter area usually visited three or four months each year. Mechanitis spp. and Hypothy- 
ris euclea leucania (Bates) were the most frequently observed ithomiines exhibiting this 
behavior. These ithomiines are very abundant, relative to others, at this locality (Young, 
1976, Pan Pacific Entomol. 53:104-113; Young, 1979, J. Lepid. Soc. 33:68-69; Young 
and Moffett, 1979a, Amer. Midl. Natur. 101:309-319; 1979b, Deutschen Ent. Zeitschr. 
26:21-38). A less numerous ithomiine, Godyris zavelata caesiopicta (Niepett) was also 
observed feeding on fresh bird droppings at various times in the same period (Fig. 1). 
In my experience, encounters of such behavior consisted of usually one or two butterflies, 
either both on the same dropping or on separate droppings in the case of two or more. 
Large aggregates of ithomiines on bird droppings were not encountered. At the same 
times, however, I did not notice any swarms of army ants in the same areas, or in adjacent 
open areas such as a cacao plantation forming the border to the forest study site. In one 
instance with Godyris (10 July 1982 at 1600 h) I noticed a single butterfly feeding at a 
dropping for close to forty minutes but with frequent interruptions by several flies (Dip- 
tera) that chased it away temporarily. Godyris zavelata females are easily distinguished 
from males by wing colors (Young, 1974, Entomol. News 85:227-238). It is by no means 
as abundant locally (in this area) as Mechanitis and Hypothyris. Several other bluish 
clear-wing ithomiines (undetermined) also visited fresh bird droppings in the same forest 
patch. 

Based upon these preliminary observations made at irregular intervals over several 
years at the same forest patch in northeastern Costa Rica, I suggest that the females of 
several genera of ithomiine butterflies routinely exploit, on an opportunistic basis, fresh 


VOLUME 38, NUMBER 1 63 


bird droppings splashed on understory vegetation. Areas of tropical rain forest with 
disruptions in the canopy, such as light gaps and foot paths, are particularly attractive 
gathering places for various species of birds, perhaps because many insects, potential 
prey, and other arthropods are also found in these microhabitats. In turn, bird droppings 
occur there frequently, although perhaps in an unpredictable fashion, selecting for op- 
portunistic foraging by female ithomiines. When large concentrations of bird droppings 
become available, such ithomiines, at least Mechanitis and Melinaea, may exhibit delib- 
erate orientation to such food resources and become abundant there, as reported else- 
where (Ray and Andrews, op. cit.). 

I thank Luis Poveda for identification of the Godyris larval food plant, and Dr. J. 
Robert Hunter for allowing access to Finca La Tigra. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 63-64 


SATYRIUM KINGI (LYCAENIDAE) TAKEN IN MARYLAND 


At 1600 h on 22 July 1982, after spending a discouraging time collecting in three areas 
in Wicomico and Worcester Counties in Maryland, I caught a Satyrium kingi (Klots and 
Clench) near Millville, Worcester County. This capture represents a significant northward 
extension of the known range of this species on the coastal plain. 

The orange cap on the blue spot on the hindwing ventrum showed the identity of this 
rare find. Its abdomen was thin, and its long tails were gone, but the slight roundness of 
its wings and the fact that it landed on a sweetgum sapling at about 5-6 feet above the 
ground corresponded with the description of Gatrelle (1974, J. Lepid. Soc. 28:33-37) of 
the flight habits of females. Its flight was slow, due possibly to its age, the lateness of the 
hour, or the deep shade in the area, but it does agree with the “sluggish” adjective used 
by Covell and Straley (1973, J. Lepid. Soc. 27:144-154). The very late date and the 
condition of the specimen (Fig. 1) indicated that this was possibly the last survivor of 
the season’s brood. 





Fic. 1. Left: S. kingi, male. Suffolk, Nansemond County, Virginia, July 1, 1974, lower 
aspect; Right: S. kingi, female, Millville, Worcester County, Maryland, July 22, 1982, 
lower aspect. 


64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The specimen was taken along a damp trail from a sandy road, where the vegetation 
consisted primarily of sweetgum (Liquidambar styraciflua L.), red maple (Acer rubrum 
L.), white cedar (Chamaecyparis thyoides (L.), loblolly pine (Pinus taeda L.), sweet bay 
(Magnolia virginiana L.), tassel-white (Itea virginica L.), blueberry (Vaccinium sp.), 
and sweet pepperbush (Clethra alnifolia L.), which was just coming into bloom. This 
habitat resembles in some respects that designated as Group A for kingi by Gatrelle 
(1974). 

Only three worn Megisto cymela (Cramer) and one fresh male Wallengrenia otho (J. 
E. Smith) were seen in the same area on the date of capture. Incisalia henrici (Grote & 
Robinson) was common and I. augustus (Kirby) rare in the previous spring, the only 
other time I had collected there. 


WILLIAM A. ANDERSEN (M.D.), 220 Melanchton Avenue, Lutherville, Maryland 21098. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 64 


THE IDENTITY OF WING HAIRS IN MEGALOPYGIDAE 


The wings of Megalopygidae were described as being ee with long, wrinkled or 
wavy hairs that gave them a wooly appearance. 

By making transparent impressions of the upper surface of the front wings of both 
male and female Megalopyge opercularis (J. E. Smith), using the replica method de- 
scribed by Khalaf (1980, Fla. Entomol. 63(3):307-340), it became clear that the wings 
were covered with scales that were deeply divided (Figs. 1 & 2); the apices were atten- 
uate; and the branches formed the so-called “hairs”. The base of the scales was cuneate 
(attenuate) as in other moths. 

This investigation received support from the Academic Grant Fund of Loyola Uni- 
versity. 


KAMEL T. KHALAF, Loyola University, New Orleans, Louisiana 70118. 





Fics. 1 & 2. Light micrograph of replica of the front wing of Megalopyge opercularis 
(J. E. Smith), showing deeply divided scales: 1, female; 2, male. 


VOLUME 38, NUMBER 1 65 


Journal of the Lepidopterists’ Society 
38(1), 1984, 65 


POPULATION OUTBREAK OF PANDORA MOTHS 
(COLORADIA PANDORA BLAKE) ON THE KAIBAB PLATEAU, 
ARIZONA (SATURNIIDAE) 


The pandora moth (Coloradia pandora Blake) is fairly widespread in the pine forests 
of the Rocky Mountains, and occasionally exhibits large population outbreaks as noted 
by Ferguson (1971. Moths of America North of Mexico, Fascicle 20.2A, E. W. Classey, 
Ltd., London). Such an impressive outbreak was noted on a visit to the Kaibab Plateau 
of northern Arizona in August 1982. During a field trip to the plateau, thousands of adult 
pandora moths were observed flying about or landed upon tree trunks in yellow pine 
(Pinus ponderosa) forest in the daytime hours. While driving a northsouth transect the 
full length of the Kaibab Plateau on 15 August 1982, the greatest concentrations of 
pandora moths were noted within a two-three mile zone surrounding the Jacob Lake 
Junction, on State Highway 89 (Alt.) and Highway 67. Hundreds of adult moths (many 
freshly emerged) and thousands of eggs were noted on the buildings and tree trunks at 
Jacob Lake, especially near outside lights that were kept on at night. 

Adult males and females were active in large numbers nocturnally as well as diurnally, 
because “black lighting” at night produced heavy catches near the North Rim of the 
Grand Canyon on 16 August. Wygant (1941. Jour. Econ. Entomol. 34(5):697-702) noted 
in Colorado that the peak emergence of adults was in July, every-other-year, because of 
a two-year life cycle, and the primary food plant was lodgepole pine (Pinus contorta). 
In another area, Oregon, yellow pine was reported to be the principal food plant of the 
pandora moth by Packard (1914. Mem. Nat'l. Acad. Sci. 12:1—276). Since the yellow pine 
predominates on the Kaibab where pandora moths were observed to be most abundant 
in August 1982, this pine is probably the most important food plant there. 

Several hundred eggs were oviposited by freshly collected females placed in glassine 
envelopes. The ova were glossy blue-green spheres which hatched in early September 
three to four weeks after oviposition. This fits with Ferguson’s notation that the young 
larvae overwinter, mostly in the second instar, on the pine branches at the base of needles. 
Attempts to rear the larvae on Pinus palustris (which was available to the author) failed. 

Adult pandora moths are clearly strong flyers, since one was observed flying across a 
barren desert landscape some 45 miles west of the edge of the Kaibab Plateau and the 
nearest pine trees. Undoubtedly, during large population outbreaks, some individuals 
wander great distances in search of suitable food plants to oviposit upon. 


LARRY N. BROWN, Department of Biology, University of South Florida, Tampa, 
Florida 33612. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 65-66 


TWO LARGE COLLECTIONS OF MACROLEPIDOPTERA 
TO THE MILWAUKEE PUBLIC MUSEUM 


The Milwaukee Public Museum in recent years has received two major Lepidoptera 
collections, the William E. Sieker Collection of Sphingidae and the James R. Neidhoefer 
Collection of Macrolepidoptera of several families. 

A donation from the wife of the late Mr. Sieker and daughter Marie, the Sieker 
Collection was acquired by the Milwaukee Public Museum in September 1982. Amassed 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


over almost fifty years, this outstanding collection totals over 9000 prepared Sphingidae 
representing some 150 genera, 1000 specific and subspecific taxa, and includes some type 
material. All major faunistic regions are well represented. Mr. Sieker acquired a major 
portion of the collection through exchange and donations from biologists conducting field 
research in different parts of the world. The collection includes all of the known Wis- 
consin sphingid species, the result of Mr. Sieker’s own collecting and his strong association 
with other naturalists in the state over the years. There are also several hundred prepared 
Catocala moths (Noctuidae), an additional several thousand papered specimens, associ- 
ated field notes, several hundred reprints of important works on sphingids and other 
groups, and reference books. 

William E. Sieker was born in 1912 in Milwaukee, Wisconsin and died in Madison, 
Wisconsin in January 1982 at age 70. Although a tax attorney by profession, he pursued 
a second career of collecting and studying sphinx moths, particularly in the northern 
reaches of Wisconsin. His interest in Sphingidae was fused with a dedication of helping 
conservation efforts, particularly in Wisconsin. Mr. Sieker was a founder of the Wisconsin 
Entomological Society and a past president of that organization as well as of the Madison 
Audubon Society. He was also legal counsel for the Wisconsin Chapter of Nature Con- 
servancy and helped that organization acquire the Ridges Sanctuary at Baileys Harbor 
in Door County. 

The museum acquired the James R. Neidhoefer Collection in January 1976, a collec- 
tion which includes about 95,000 specimens of Macrolepidoptera (approx. 45,000 pre- 
pared and 50,000 papered specimens) of which approximately 1200 are gynandromorphs, 
sexual mosaics and aberrations (structural and color). The collection is particularly strong 
in Papilionidae, Nymphalidae, Heliconiidae, Ithomiidae, Morphidae, Pieridae, Saturni- 
idae and Sphingidae. All of the major faunistic regions are represented, with particular 
strengths in the Neotropical and Indo-Australian Regions. Mr. Neidhoefer’s donation also 
included 182 insect storage cabinets with 105 drawers, and an extensive library of rare 
books, reprints, and monographs on the Lepidoptera. At the time the collection was 
donated to the museum, Mr. Neidhoefer also financed the renovation of a collection 
storage room in the Invertebrate Zoology Section (which includes entomology). 

Mr. Neidhoefer acquired his collection over forty years, through buying and exchang- 
ing specimens with collectors all over the world, and through field expeditions (i.e., Brazil) 
financed by him and for the purpose of collecting. With the cooperation of a former 
curator of the Milwaukee Public Museum, Kenneth MacArthur, Mr. Neidhoefer was 
instrumental in acquiring other Lepidoptera collections for the museum, most notably 
the George Berg Collection (which includes a good series of Nicaraguan Rhopalocera), 
the P. Gagarin Collection (Brazil), and others. 

James R. Neidhoefer was born in Milwaukee in 1917 and became an avid naturalist 
at an early age. He received an undergraduate degree in zoology from Marquette Uni- 
versity and did a thesis on the freshwater sponges of Wisconsin. He took over the family 
carpet business in Milwaukee but continued to pursue his interests in natural history by 
collecting Lepidoptera and teaching his 12 children about insects. Prior to moving to 
Miami, Florida in 1981 to pursue a new retirement job as president of a wholesale pet 
distributorship, Mr. Neidhoefer was very active in local nature organizations and the Boy 
Scouts, as well as the Milwaukee Public Museum. As an honorary curator for the museum, 
he now collects Lepidoptera and other invertebrates in Florida during his spare time. 

With the acquisition of the Neidhoefer Collection, and through grants from the Insti- 
tute of Museum Services and the Friends of the Milwaukee Public Museum, a major 
collection reorganization and upgrading of facilities was initiated by the museum’s full 
time curators in the Lepidoptera area, myself and Susan S. Borkin. With the acquisition 
of the Sieker Collection, and combined with further collecting efforts in Wisconsin and 
also from ecological studies in the Neotropical Region, one of our goals is to make these 
outstanding collections of use to curators, systematists, and biologists working on groups 
represented in them. 


ALLEN M. YOUNG, Curator and Head, Invertebrate Zoology Section, Milwaukee Pub- 
lic Museum, Milwaukee, Wisconsin 532338. 


VOLUME 38, NUMBER 1 67 


Journal of the Lepidopterists’ Society 
38(1), 1984, 67 


ARE CHAIN-LINK FENCES BARRIERS TO BUTTERFLIES? 


During the summers of 1982 and 1988, I regularly collected European cabbage but- 
terflies, Pieris rapae Linnaeus from the Fenway Victory Gardens, Boston, Massachusetts, 
and Dunback Meadows, Lexington, Massachusetts. From 24 June to 18 August 1982, and 
2 June to 1 August 1983, I observed 27 confrontations between free flying P. rapae and 
chain-link fences. On each occasion the butterfly flew within 5-10 cm of the fence, back 
and forth over a 1 to 1.5 meter area, and then added a vertical movement of equal 
distance. Three times P. rapae succeeded in flying over the fence. Once a male flew to 
the end of the fence and around if, and once a butterfly proceeded after a 2-3 second 
delay to pass through the fence after I tried unsuccessfully to capture it. On 21 occasions 
P. rapae changed their flight direction nearly 180° after confronting chain-link fences. 
On one occasion an alfalfa butterfly, Colias eurytheme Bdy. was observed to change 
direction approximately 90° after confronting a fence. A 90° change was also observed 
once for a P. rapae after physically striking a fence. The openings in a chain-link fence 
measure approximately 7 cm in height and width. The mean wing spread of P. rapae is 
only 3.8 cm. On several occasions I have seen individual P. rapae squeeze their folded 
wings through 1.8 cm wire screening of a flight cage in the laboratory; and in the field, 
I have observed individuals fly without hesitation through thin wire fences with openings 
of 12-15 cm. Even though chain-link fences have openings through which a P. rapae 
could physically pass without contact, the butterfly rarely does so. Perhaps P. rapae can 
not accurately judge the opening size; it may appear small and likely to damage wing 
tips; or perhaps the thick shiny wire on all sides of the butterfly may be distorted by the 
butterfly’s visual system and perceived as a nearly solid barrier. 

Chain-link fencing is used widely to keep would-be intruders out of areas or keep in 
desired objects. Mountain alpine areas are under increasing pressures from human visitors 
each summer. Some parks have posted personnel to keep visitors on established trails, 
others have begun to rope off areas. Chain-link fences have been proposed as a means 
to save badly trampled alpine areas. 

The construction of chain-link fences and other obstacles may have a variety of effects 
on butterfly populations depending on the species involved and the habitat. Williams 
(1930. The migration of Butterflies, Oliver and Boyd, London. 473 pp.) states that Be- 
lenois severina and Vanessa cardui usually fly over obstacles with little or no lateral 
deviation from their line of flight. Feltwell (1982. Large White Butterfly The Biology, 
Biochemistry, and Physiology of Pieris brassicae (Linnaeus), Dr. W. Junk Publishers, 
The Hague, 535 pp.) reports that P. brassicae typically flies over obstacles rather than 
around them. However, Andronymus neander predominantly flies laterally with little 
or no vertical rise when confronted by an obstacle in its flight path (Williams, 1930. 
ibid). Generally, alpine lepidoptera fly very low to the ground to avoid winds. If fences 
are encountered, movement may be hindered, adding an additional energetic pressure 
on mountain butterfly populations which are often already low in number. Therefore, 
there may be serious deleterious effects on alpine butterfly populations if chain-link 
fences are built in these areas. 

These observations are limited in number and species involved. Perhaps a more quan- 
tified investigation is merited. Such an investigation should be concerned with the height 
and opening sizes of fences, with a look at a number of different species in various 
habitats to determine if the observations reported here can be generalized. 


MARK K. WourMs, Department of Biology, Boston University, 2 Cummington Street, 
Boston, Massachusetts 02215. 


Journal of the Lepidopterists’ Society 
38(1), 1984, 68 


BOOK REVIEW 


CATALOGO SISTEMATICO DE LOS LEPIDOPTEROS IBERICA. (I) MACROLEPIDOPTERA, by M. 
R. Gomez-Bustillo and M. Arroyo-Varela. 1981. Inst. Nac. Invest. Agrarias, Ministerio de 
Agricultura y Pesca, Madrid. 499 pp., 6 col. pls. (1200 Pta. [=$9.40)). 


Recent catalogs and checklists, including those of Bradley et al. (1972) for England, 
Karsholt and Nielsen (1976) for Denmark, and Leraut (1980) for France and Belgium, 
have almost covered the entire Lepidoptera fauna of the most western parts of Europe 
with up-to-date checklists. The new catalog by Gomez-Bustillo and Arroyo-Varela closes 
the gap by covering the fauna of Spain and Portugal. Their work is the first of two 
volumes; the second volume is to cover the Microlepidoptera. The catalog initially strikes 
one as very different from most catalogs, since the cover has a large color photograph of 
the pierid Aporia crataegi (L.), not what one usually finds on catalog covers. Additionally, 
there are six color plates near the back of the book with photographs from nature of a 
representative species of each family in the book. The text is also untraditional inasmuch 
as bibliographic references are included for each family in terms of literature on the 
species of the Iberian Peninsula. There is an initial brief summary of the classification 
adopted for the catalog, generally following recent classifications like that of Common 
(1970, Insects of Australia), followed by a short introduction on the origins and evolution 
of Iberian Lepidoptera. The main text treats 39 families of so-called Macrolepidoptera, 
grouping many primitive families together with the normal macros. This arrangement 
produces an artificial and utilitarian arrangement for the catalog designed to conform to 
the older concepts, whereby large-sized moths were placed in “Bombyces’’. This is not 
altogether detrimental, since a phylogenetic chart of families is included, but it does 
maintain the myth that these “Bombyces” are somehow related more than they really 
are, and it also detracts from a strictly systematic treatment of families from primitive 
to more advanced. Nonetheless, the catalog is a welcome addition to the works listing 
the European fauna. 

The authors follow a family usage that splits families too much, in my view, but does 
follow the practice of many European specialists. Thus, such groups as Syssphingidae, 
Riodinidae, Danaidae, Thaumetopoeidae, Dilobidae, and Ctenuchidae, which many con- 
sider only of subfamily status, are here raised to family level. I did not make any detailed 
checks of nomenclature. In Sesiidae, however, not all synonymies are included for each 
species, only a few of the major ones. Each family name is provided with authorship 
and dates, as well as for other higher categories. The specific and generic checklist then 
follows, with a discussion section and reference list for each family. The species are all 
listed with their dates of authorship, with parentheses added when the names of species 
have been recombined. The names of subspecies and forms, however, are not provided 
with dates. Each species is also given a notation as to its place in a European faunal 
district; thus, statements are made such as “endemic to Iberia” or “supramediterranean. ” 

The catalog is in spanish, but since the main text involves a checklist of taxa of the 
Iberian Peninsula, it is easily used by anyone. It is a welcome addition to the growing 
rank of faunal catalogs and checklists. One can only hope that now, in lieu of a new 
checklist of the entire Palearctic region, others will follow the lead of the authors and 
provide additional regional catalogs (e.g., the Balkans, Russia, the Far East), so that in 
this way we may in time have new lists of all the areas within the Palearctic region. 


J. B. HEPPNER, Department of Entomology, Smithsonian Institution, Washington, 
D.C. 20560. 


Date of Issue (Vol. 38, No. 1): 27 July 1984 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MacpDa R. Papp, Editorial Assistant 
Douc.Las C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 
NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name, with authors of 
zoological names. Insect measurements should be given in metric units; times should be 
given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where 
italics are intended. References to footnotes should be numbered consecutively, and the 
footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

Illustrations: All photographs and drawings should be mounted on stiff, white back- 
ing, arranged in the desired format, allowing (with particular regard to lettering) for 
reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches 
are not acceptable and should be reduced photographically to that size or smaller. The 
author's name, figure numbers as cited in the text, and an indication of the article's title 
should be printed on the back of each mounted plate. Figures, both line drawings and 
halftones (photographs), should be numbered consecutively in Arabic numerals. The term 
“plate” should not be employed. Figure legends must be typewritten, double-spaced, on 
a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, 
with a separate paragraph devoted to each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material should be kept to a minimum and 
must be typed on separate sheets, and placed following the main text, with the approx- 
imate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer's errors. Excessive author’s changes at this time will be charged to 
authors at the rate of 75¢ per line. A purchase order for reprints will accompany the 
proofs. 

Correspondence: Address all matters relating to the Journal to the editor. Short 
manuscripts such as new state records, current events, and notices should be sent to the 
editor of the News: June Preston, 832 Sunset Drive, Lawrence, Kansas 66044 U.S.A. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


THE LIFE HISTORY AND ECOLOGY OF EUPHYDRYAS GILLETTII 
BARNES (NYMPHALIDAE). Ernest H. Williams, Cheryl E. 
Holdren & Paul R. Ehrlich ee 

CORRECT NAME FOR THE NEOTROPICAL SQUASH-VINE BORER (SE- 
SUDAE: MELITTIA). Vitor O. Becker & Thomas D. Eichlin 


LIFE HISTORIES OF FOUR SPECIES OF PHILIRIS ROBER (LEPI- 
DOPTERA: LYCAENIDAE) FROM PAPUA NEW GUINEA. 
Michael Parsons 22 ee 


COURTSHIP BEHAVIOR OF THE GULF FRITILLARY, AGRAULIS VA- 
NILLAE (NYMPHALIDAE). Ronald L. Rutowski G John 
Schaefer 00 Ee 


CHECKLIST OF MANITOBA BUTTERFLIES (RHOPALOCERA). Paul 
Klassen 2.00 ee ee ee 


THE LIFE HISTORY AND BEHAVIOR OF EPIMARTYRIA PARDELLA 
(MICROPTERIGIDAE). Paul M. Tuskes & Norman J. Smith 


A NEW ACANTHOPTEROCTETES FROM THE NORTHWESTERN 
UNITED STATES (ACANTHOPTEROCTETIDAE). Donald R. 
DGbis a ee 


Two INTERESTING ARTIFICIAL HYBRID CROSSES IN THE GENERA 
HEMILEUCA AND ANISOTA (SATURNIIDAE). Richard Steven 
Peigler & Benjamin D; Williams _.... ee 


SPERMATOPHORE PERSISTENCE AND MATING DETERMINATION IN 
THE Gypsy MOTH (LYMANTRIIDAE). Cynthia R. Loerch & 
E. Alan: Camerore 2c 


GENERAL NOTES 


Insect parasites and predators of hackberry butterflies (Nymphalidae: Aster- 
ocampa).. Timothy P. Friedlander: 2.0304 2) J 


Ithomiine butterflies associated with non-antbird droppings in Costa Rican 
tropical rain forest. Allen M. Young 201000)» 


Satyrium kingi (Lycaenidae) taken in Maryland. William A. Andersen .. 
The identity of wing hairs in Megalopygidae. Kamel T. Khalaf |... 
Population outbreak of pandora moths (Coloradia pandora Blake) on the 

Kaibab Plateau, Arizona (Saturniidae). Larry N. Brow? .-cccccccsccceeneeneee 


Two large collections of Macrolepidoptera to the Milwaukee Public Mu- 
seum. Allen M. Young 


Are chain-link fences barriers to butterflies? Mark K. Wourms 


BOOK REVIEW 09.2500 Cl eS 


13 


15 


23 


32 


40 


47 


ol 


O7 


1984 Number 2 


Volume 38 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


Published quarterly 


Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


tee ee 


. SA 
. = { 

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——_—_ Ss 
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16 August 1984 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


LEE D. MILLER, President CHARLES V. COVELL, JR., 
KAROLIS BAGDONAS, Vice President Immediate Past President 
MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary 
J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


K. S. BROWN, JR. F. S. CHEW J. M. BURNS 
E. D. CASHATT G. J. HARJES F. W. PRESTON 
T. C. EMMEL E. H. METZLER N. E. STAMP 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229, U.S.A.; and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- 
temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- 
sion in the Sesiidae fascicle for the Moths of America North of Mexico. The dusky 
clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of 
willows and poplars. 


JOURNAL OF 


Tue LeprporreRrists’ SOCIETY 











Volume 38 ~ 1984 Number 2 


Journal of the Lepidopterists’ Society 
38(2), 1984, 69-84 


LIFE HISTORIES OF TAENARIS (NYMPHALIDAE) 
FROM PAPUA NEW GUINEA 


MICHAEL PARSONS 


Insect Farming & Trading Agency, Division of Wildlife, 
P.O. Box 129, Bulolo, Morobe Province, Papua New Guinea 


ABSTRACT. Descriptions and illustrations of the early stages and ecology of Taen- 
aris onolaus Kirsch and Taenaris catops Westwood are given with a brief description 
and illustrations of the early stages of Taenaris myops Felder. Adults of both T. onolaus 
and T. catops were frequently seen imbibing cycad juices which probably enhances their 
assumed distastefulness to predators. Their foodplant specializations and aposematic at- 
tributes are discussed together with the mimetic relationships of Taenaris. 


The genus Taenaris Hiibner in Papua New Guinea numbers 18 
species. Together with three species of the genus Morphopsis Oberthiir 
and the monotypic genera Hyantis Hewitson and Morphotenaris 
Fruhstorfer, these are the only representatives of the Morphinae to be 
found in the country. A further six species of Taenaris and one of 
Morphopsis are known from Irian Jaya. Torres Strait marks the bound- 
ary of the distribution of these few closely related genera and species 
in the Melanesian region. They do not occur on the Australian main- 
land. 

The Morphinae occur widely throughout the Indo-Australian region 
and number about 100 species. The morphology of the early stages 
and of the adults indicate their close affinity with the Satyrinae. For 
example, adults of Morphopsis albertisi Oberthiir in Papua New Guinea 
superficially resemble the smaller satyrine Tisiphone helena Ollift. 
from north Queensland, Australia, where no mimicry could be in- 
volved. 

Adults exhibit little sexual dimorphism, but males tend to be smaller 
than the females, have a more concave inner margin of the forewing 
and bear sub-basal androconial tufts on the hindwing. 

Little was known of the biology of the Morphinae in the Melanesian 
region. Rosier (1940) gave some details of the biology of Taenaris 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


horsfieldii Swains. from Java and D’Abrera (1977) mentioned briefly 
the early stages and foodplants of Taenaris catops Westwood and 
Taenaris phorcas Westwood. D’Abrera stated that a paper describing 
the life history of M. albertisi was in preparation, but until now there 
has been no detailed study of any species of this subfamily published 
for the region. 


Taenaris onolaus Kirsch 


D’Abrera (1977) lists four races of this species, two occurring in Irian 
Jaya. The description of the subspecies ida Honrath fits the butterfly 
described here and this, therefore, represents an extension of its range 
from the known type localities for the subspecies in the Huon Penin- 
sula. The 10 km grid square reference in which the subspecies has been 
found in Bulolo is DN50 at approximately 700 m. The present study 
was made from October to December 1979. 


Egg (Fig. 2). 1.5 mm in diameter; pearly white when laid, changing within two days 
through cream to deep pink; almost spherical, but slightly tapered towards flattened 
apex; chorion covered with evenly spaced, shallow dimples. Duration, 14 days. 

Larva. First instar. Length 4 mm on hatching, 5 mm at end of instar; head jet black, 
shiny with fine white setae; thorax and abdomen with fine white setae up to 1 mm in 
length, initially cream, gradually changing to yellowish green, then orange-red; prothorax 
with dorsolateral black spots. Duration, 2 days, and a further 5 days of inactivity during 
pre-ecdysis and ecdysis. 

Second instar. Length 10 mm at end of instar; head jet black, shiny, 1 mm in diameter 
with setae 3.5 mm in length and pair of truncate, slightly forwardly curved horns 0.75 
mm in length, each horn with 3 strong spines; thoracic setae 4 mm, abdomen with setae 
3.5 mm in length; thorax and abdomen deep pink, abdomen with a dorsal black spot on 
anal segment. Duration, 4 days, plus 3 days of inactivity during pre-ecdysis and ecdysis. 

Third instar. Length 22 mm at end of instar; similar to second but head 2 mm in 
diameter, horns 1 mm in length, each with 5 spines; thoracic setae 6 mm; thorax and 
abdomen pink with 4 indistinct, but continuous orange-yellow lines, 2 dorsolateral and 
2 lateral. Duration, 6 days, plus 2 days of inactivity during pre-ecdysis and ecdysis. 

Fourth instar. Length 35 mm at end of instar; head 3 mm in diameter, horns 1.5 mm, 
each with 6 spines; thorax and abdomen wine-red, orange-yellow lines slightly more 
prominent; body setae up to 9 mm in length; below these a layer of strong, sharp, black 
setae 1.55 mm in length. Duration, 5 days, plus 2 days of inactivity during pre-ecdysis 
and ecdysis. 

Fifth instar (Fig. 4). Length 60 mm at end of instar; similar to fourth but head 5 mm 
in diameter, horns 2.5 mm with 6 strong spines (Fig. 20c); body setae up to 10 mm; 
lower black setae 3 mm. Duration, 8 days, plus 2-3 days spent wandering. 

Prepupa. Larval color changes from wine-red to yellow after suspension prior to 
pupation so that lower black setae and black spots of prothoracic and anal segments 
become very prominent. Duration, about 1 day of hanging before larva to pupa ecdysis. 

Pupa (Figs. 6 & 7). Length 30 mm; ovate, smooth, translucent creamy white; cremaster 
black; anal rise with 2 black tubercles; apical margin of front bifid, forming 2 short, 
conical horns above each eye 1 mm in length. Duration, 17-20 days. 


Ecological Observations 


Foodplant and habitat. The foodplant is, unusually, a gymnosperm, 
Cycas circinalis (L.) Laut. & K. Sch. of the order Cycadales. This, 


VOLUME 388, NUMBER 2 Teak 





Fics. 1-9. Taenaris onolaus: 1, female ovipositing on cycad; 2, eggs; 3, second instar 
larvae; 4, mature larva; 5, mature larvae at rest in the leaf litter near the base of their 
foodplant; 6, dorsal profile of the pupa; 7, lateral profile of the pupa; 8, upperside of 
male; 9, underside of female. 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


however, is not a unique specialization among the butterflies. Taenaris 
butleri Oberthiir is known from the same foodplant (T. Fenner, pers. 
comm.) and also Luthrodes cleotas Guérin of the Lycaenidae (Szent- 
Ivany et al., 1956). Another lycaenid, Theclinesthes onycha Hewitson, 
is also known from Cycas in Australia (Sibatani & Grund, 1978). 

The foodplant exhibits a well defined distribution in the study area, 
being restricted to a well drained ridge alongside a gravel road behind 
the Bulolo Forestry College. The cycads cover about 3 acres which 
makes the area ideal for the study of a defined population of T. ono- 
laus. 

Cycas circinalis occurs locally at a quite high density (in places up 
to six plants per 10 square meters), mainly along the top of the ridge 
and under a 15 year old Pinus plantation. The plantation provides a 
fairly open habitat with only semi-shading by the thin pine canopy. 
Saplings of other trees occur sporadically throughout the plantation. 
These conditions appear to be ideal for the growth of the cycads and 
may explain why the plant is not found locally outside the area where 
the scrub becomes thicker. A number of plants were fruiting prolifi- 
cally during visits to the area in October, November and December 
1979, and there were many cycad nuts on the ground. 

Oviposition and phenology. Eggs are laid by females in batches 
ranging in number from 20 to 40 with an average batch size of about 
30. The highest number recorded in a single batch was 77. They are 
deposited close together, but not touching, on the undersides of one 
(or sometimes two) leaves of the older, tougher, dark green fronds. 
They are always placed about one third of the way down from the tip 
of the frond. Occasionally (seen in at least seven batches), there are 
one or two unfertilized eggs which remain white after the others have 
changed to pink. 

The plants on which the females choose to oviposit are all of about 
the same height, approximately 1.5 m tall, and with usually 5-15 
fronds. Cycads are extremely slow growing, and these plants are esti- 
mated to be from 5-6 years old (possibly older). As yet they have little 
or no trunk, and the fronds of most of them arise directly from the 
ground. No eggs or larvae were found on the younger plants with only 
two or three fronds and of smaller overall size at the beginning of the 
study period. 

From observations of two females made late one afternoon in De- 
cember from 1735 h onwards, it appears that T. onolaus only oviposits 
during the period of about two hours before complete darkness which 
is at 1900 h, dusk (or half-light) coming at about 1830 h. (This was 
suggested later by two further observations of females ovipositing at 


VOLUME 388, NUMBER 2 73 


dusk.) One female was discovered at 1745 h below a cycad frond, 
having laid about 30 eggs. Approximately every two minutes she de- 
posited another egg in the row of four across the cycad leaf. Having 
completed a row she then moved slowly forward and positioned herself 
to begin a new row, from side to side. It is estimated, therefore, that 
a whole batch of about 50 eggs (this particular female had gone by the 
next morning but laid 45 eggs) would take approximately two hours 
to lay. 

A second female was seen at the same time flying around another 
cycad, repeatedly settling on the upperside of a frond and then crawl- 
ing beneath it. She then flew behind some vegetation, which obscured 
the other half of the same cycad, and settled out of sight. Soon after 
she was re-located sitting on a batch of about 25 newly laid eggs ready 
to recommence egg laying. It appears, therefore, that some females 
take periods of rest away from the cycads on which they are ovipositing 
then return to lay their eggs at intervals. Both females were still ovi- 
positing in near darkness at 1850 h. 

It is possible that females are able to lay further batches of eggs. 
However, it appears that their ovaries produce a certain number of 
eggs that are laid as a single batch in a short period of time. They are 
probably fairly short lived once they have paired and have finished 
ovipositing. It is also evident that females can detect the presence of 
eggs or larvae (probably visually) that are already present on the cycad, 
because when the area was studied in mid-November, no suitable look- 
ing plant was found to have more than one batch of eggs or larvae on 
HU 

Eggs of T. onolaus were first discovered at the beginning of October 
1979 which marked the end of an extremely dry dry-season. This lasted 
from the end of June for three months. During this time there was no 
rain recorded for the Bulolo Valley. In October, however, there were 
a few batches of T. onolaus larvae to be found, mainly fourth and fifth 
instars. Two egg batches were located at this time, which indicates 
that, even though the climate can be seasonally extremely dry in the 
area, generations can be continuous throughout the year because of 
the hardiness of the cycads; their foliage remains constant all the year 
round. (During the 1979 dry season in the Bulolo Valley, many angio- 
sperms, especially vines, even in fairly dense forest areas, began to wilt 
and/or ceased new leaf growth. Often the dry season is hardly appar- 
ent.) 

When the area was again revisited at the end of the first week in 
November, heavy rains had recommenced during the previous four 
weeks. At this time all but about eight suitable-looking plants had eggs 


74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


or larvae, and the approximate census was as follows: 18 batches of 
eggs, 7 groups of first instar larvae, 5 groups of second instar larvae 
and 2 of third instar larvae. 

During the latter half of December females were still ovipositing, 
and a number of cycads, even the smaller, single-frond plants, were 
seen with newly laid egg batches. Some hosted up to three age classes 
of larvae, all of which fed together. This suggested that the area was 
now almost saturated with early stages due to the build up in numbers 
of adults and that intraspecific competition can occur where the species 
has limited, or finite, food resources. The few suitable cycads with no 
larvae at this time implied that any eggs laid on them may have been 
the subject of predation. Although no predators have been seen taking 
early stages, certain fresh egg batches were often found to have been 
eaten by the next day. Tetigoniid grasshoppers and predatory bugs are 
likely to be responsible. 

In general, it may be concluded that as T. onolaus is a cycad feeder 
of tropical distribution, it is subject to little periodicity, i.e., that it is 
continuously brooded all year round, but that any large fluctuation in 
population numbers is reciprocal of extremes of wet and dry weather. 
Prolonged dry periods appear to produce aestivative (diapause) pupae 
and may also retard growth of new cycad fronds, so that the result at 
the onset of new rains is a large buildup of adults and early stages 
which compete intraspecifically for foodplants in areas with limited 
distribution of cycads. 

Larval behavior. Larvae of T. onolaus are gregarious throughout 
their feeding period. In the fourth and fifth instars, however, the dis- 
tance that separates each larva is increased, and they may be found 
feeding singly, or in sub-groups of up to five. First to third instar larvae 
spin an almost invisible mat of extremely fine silk on which they rest 
below the frond of the cycad, so that when the plant is viewed from 
above they are completely obscured from the observer. 

When feeding, early instar larvae begin at the tips of the leaves of 
the cycad frond and eat each leaf back separately to the base of the 
main stem. The group will then begin to feed again on the next leaf 
and progress gradually downwards. They often defoliate a whole frond 
as they grow. The smaller larvae form very orderly rows when resting 
or as they feed on the edge of the leaf lamina (Fig. 3). Final instar 
larvae tend to be cannibalistic on soft, newly formed pupae if many 
are caged together. One particular batch of about 12 fourth instar 
larvae were found resting during the day at the base of a frond of one 
cycad and were thus hardly visible beneath the leaf litter trapped there 
(Fig. 5). This does not, however, appear to be typical behavior. They 
were not undergoing ecdysis, and it is possible that these larvae were 


VOLUME 38, NUMBER 2 ‘ko 


feeding at night and seeking shelter from predators during the day. 
All instars have been found feeding at various times during the day 
with no specific feeding or resting times. 

In general, all instars are fairly slow in their movements. When 
touched they sometimes react by thrashing the head from side to side. 
This appears to be an effective means of warding off insect predators. 
Fifth instar larvae tend to curl up and fall off the cycad fronds if 
handled, behavior which enhances their very moth-like appearance. 

Adult behavior and abundance. Females appeared to be most fre- 
quent in the study area and were seen at all times flying randomly 
throughout the pine plantation. Specimens were seen on every visit 
during the study period although never in great abundance at any one 
time. Numbers ranged on average from 1-4 (flying in close proximity) 
seen per hour. 

At one time in mid-December a sample census resulted in the sight- 
ings during one hour of three males flying in a restricted gully on the 
border of the study area (one feeding on damp mud), and four females. 
Two of these females hung inertly beneath a cycad frond (late after- 
noon) and did not react to rapid hand movements nearby. One was 
picked up and promptly flew off when released. Invariably, however, 
adults are very wary and do not allow one to approach to within less 
than 2 meters if they are at rest and alert on the upperside of broad- 
leaved foliage. 

There are no succulent fruit trees in the pine plantation, and none 
of the saplings which produce small berries that fall to the ground have 
proven attractive to adults. However, on a number of occasions both 
sexes have been seen feeding on the fermenting skins of cycad nuts 
that have fallen to the ground when brown and ripe. 

Competition. Intraspecific competition has been mentioned. How- 
ever, there also exists in the area, interspecific competition for Cycas 
circinalis between T. onolaus and a chrysomelid beetle of the subfam- 
ily Criocerinae. The small 1 cm long, orange beetle which is probably 
Crioceris clarkii B. Baly (based on the discussion in Szent-Ivany et al., 
1956), feeds as a cream colored larva on the cycads. It has a definite 
preference for the soft, new, light-green cycad fronds. Therefore, by 
selecting only the older tougher fronds on which to oviposit, T. onolaus 
probably avoids competition for individual plants. Nevertheless, the 
beetle does cause much damage to the cycads in the area and can be 
classed as a successful competitor with T. onolaus. 

The feeding damage caused by the beetle larvae is very character- 
istic. Even for a long time after a cycad has been eaten back by either 
herbivore it is possible to determine whether it was fed on by beetle 
or butterfly. Whereas T. onolaus eats the whole leaf of a frond, the 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


chrysomelid eats only the underside of a lamina and leaves the top 
waxy cuticle as a window. This soon dries, turns yellow, and is left 
trailing, still attached to the frond. 

Many new, recently unfurled, cycad fronds were the subject of at- 
tack by the beetle in mid-December 1979. These beetles also appear 
to play a significant réle in controlling the population size of the but- 
terfly. It is possible that they cause a final crash in the numbers of a 
cohort of T. onolaus because, if there are sufficient numbers of the 
beetle, then growth of cycads in the area may be halted completely. 
There will not, therefore, be enough fronds which reach maturity for 
the benefit of T. onolaus. 


Taenaris catops Westwood 


D’Abrera (1977) lists 21 races of this species. As emphasized by 
Brooks (1950) the named subspecies of T. catops may be very artificial 
as the species is widely distributed in New Guinea, of common status, 
and exhibits a great phenotypic variability both locally and regionally. 
Considering for example the supposed subspecies mylaecha from Sud- 
est Island which is described by D’Abrera as an “albinotic extreme” 
(i.e., very white), the same form is now recorded widely from the 
Western Highlands Province of the mainland (Fig. 22). The other ex- 
treme is an extremely dark form of T. catops, in which black and dark 
grey have replaced almost all the white. Supposed subspecies of T. 
catops should, therefore, be accepted with caution and are more likely 
the result of clinal variation or Miillerian mimetic associations within 
their genus. 

The life history of this species was also recorded from the T. onolaus 
study area in March and April 1980. The egg and first two instars 
cannot be described as only the third instar onwards were available. 


Third instar (Figs. 10 & 11). Larvae grew extremely rapidly from 8 to 25 mm in 3 
days; head jet black, shiny, 2 mm in diameter, covered with fine white setae, horns 
similar to those of T. onolaus, 1.5 mm in length; body covered with soft, white setae, 
longest (5 mm) on the prothoracic and anal abdominal segments, decreasing to 4 mm at 
body center; thorax and abdomen dark grey with 2 dorsolateral and 2 lateral white lines; 
spiracles encircled with yellowish orange; claspers laterally yellowish orange, dorsally 
with black patch surmounted by two short (0.55 mm) pointed tubercles. Duration, 3 
days, plus a day of inactivity spent during pre-ecdysis and ecdysis. 

Fourth instar. Length 32 mm at end of instar; similar to third but body laterally black 
with middorsal black line bordered with grey. Duration, 7 days, plus 1% days of inactivity 
spent during pre-ecdysis and ecdysis. 

Fifth instar (Fig. 12). Length at end of instar 57 mm; similar to fourth but head 4.5 
mm in diameter, horns with 8 long, thin spines (Fig. 20a); body jet black but for 2 
dorsolateral white lines and 2 lateral yellow lines; spiracles black encircled with orange; 
soft white setae 7 mm longest; strong, sharp lower setae tan brown, 2.5 mm in length. 
Duration, 9 days. 

Pupa (Figs. 13 & 14). Length 31 mm; smooth ovate, pale green; cremaster pale yellow, 


VOLUME 38, NUMBER 2 eT. 


“pup 


i 


7 
Le, (i a Zz é 
Fuss AT bs ba ty ge oe 
2 eth ale e 
ee 


3 7; ee Pe 
a ries Le DP tne 6 yt ae 


x 





12 alte ee 


Fics. 10-15. Taenaris catops: 10, third instar larvae at rest; 11, third instar larvae 
feeding; 12, mature larva; 13, ventral profile of pupa; 14, lateral profile of pupa; 15, 
adult female imbibing the juices of a damaged cycad nut. 





tipped with black. In shape pupa like that of T. onolaus but frontal horns slightly longer, 
more pointed, tipped with yellow and below this is a ring of pale brown; tubercles of 
anal rise not as prominent as those of T. onolaus, only faintly tipped with brown. Du- 
ration, 12 days. 


78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ecological Observations 


Foodplant and habitat. The early stages of T. catops were discov- 
ered at the center of the T. onolaus study area previously described. 
The foodplant is a new record for T. catops. It is a 1.5 m tall ground 
orchid with large, predominantly white flowers, Phaius tancarvilleae 
(Banks in L’Herit) Bl. The plant has been found in the Bulolo Valley 
at 800 m growing under Pinus in the plantation. This may not, how- 
ever, be the usual foodplant for the species as D’Abrera (1977) states 
that T. catops feeds on the Black Palm (Caryota rumpha: Palmae), 
Betel-nut palm (Areca catechu: Palmae) and Banana (Musa: Musa- 
ceae). Pyle and Hughes (1978) list T. catops from Cordyline terminalis 
(Liliaceae) which is used in hedges in many highland areas of the 
mainland. 

Larval behavior. Like T. onolaus, the larvae are gregarious and 
remain so up to the final instar. They feed in line from the tip of the 
leaf lamina and eat the blade gradually downwards to halfway or a 
little less (Fig. 11). The larvae pause at intervals and then move slightly 
back up the blade to rest. 

Adult behavior and abundance. Females of. catops, like those of 
T. onolaus, were more often encountered in the area than males. They 
were also slightly more abundant than those of T. onolaus. Occasion- 
ally, up to five at one time were seen in one area. 

T. catops has also been found just before dusk hanging inertly be- 
neath foliage. Only at this time can they be approached because they 
are otherwise always alert and wary when feeding or resting on the 
uppersides of leaves. In forest areas they prefer to fly in shade. The 
species has been observed in many localities on the mainland flying 
just above the leaf litter in search of fermenting fruits on which to 
feed or probing moist leaf litter. 

In spite of their preference for shady habitats, both sexes of T. catops 
can commonly be seen flying through gardens in Bulolo and in straight 
lines across any open grassland areas in the Bulolo Valley. In sharp 
contrast, T. onolaus has never been observed outside the study area. 

It is interesting to note that both T. catops and T. onolaus were 
fond of visiting the fermenting husks of cycad nuts on the ground (Fig. 
15). At one time a female of T. onolaus was seen feeding between two 
T. catops females. At another time five T. catops were flushed from 
beneath two close-growing cycads on which the chrysomelid beetle 
larvae were feeding. They were seen to probe the fresh green frass of 
the beetle larvae where it had fallen to the ground. On numerous 
occasions the cut ends of cycad fronds on the ground which had exuded 
sap were seen to be extremely attractive to T. catops—this is discussed 
further below. 


VOLUME 38, NUMBER 2 79 





Fics. 16-19. Taenaris myops: 16, mature larva; 17, prepupal larva; 18, latero- 
ventral profile of pupae; 19, underside of male. 


Taenaris myops Felder 


D’Abrera (1977) lists 13 races of this species. Its full life history was 
studied from a batch of 37 eggs. These were collected from the un- 
derside of the leaf of a monocotyledon, Tapenochilus sp., of the Cos- 
taceae found in November 1980, growing on a creekside near Eilogo 
Falls (Port Moresby, Central Province, 10 km grid square EK45). This 
represents a new foodplant record. However, as the author, after lo- 
cating the eggs and watching them hatch, had other commitments, the 
early stages of T. myops were reared and photographed by Peter Clark. 
He noted that, in general, the whole life history was similar to that of 
T. catops. 

Egg. Slightly lighter pink but otherwise similar to that of T. onolaus. 

Larva. First instar. 4 mm long on hatching; head jet black, shiny, covered with fine 
white setae; body with long fine white setae, up to 1 mm in length; thorax and abdomen 
Opaque creamy white, gut from behind head to last 4 abdominal segments shows as 
pinkish red line, anal segments with traces of pinkish red. 

Second to fourth instars. Larvae at each instar exhibited similar growth rates and 
maximum sizes as those of T. catops. They grew steadily darker so that by fourth instar 


they were brownish black. 
Fifth instar. Length 59 mm at end of instar; head horns with 6 spines (Fig. 20b); 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


20 


Fic. 20. Frontal profile of left horn and lateral profiles of Taenaris final instar larval 
head capsules: a, T. catops; b, T. myops; e, T. onolaus. 


prothoracic segment wine-red, remainder of thorax and abdomen jet black, not lined as 
larvae of T. onolaus and T. catops; spiracles black, encircled with wine-red; body setae 
soft, long, white, laterally 4 mm, dorsally 10 mm in length; lower strong, black setae 4 
mm longest. 

Prepupa (Fig. 17). Larval color changed to dark grey dorsally and pale green ventrally 
after larvae had suspended themselves prior to pupation. Pupation took place 40-42 days 
after larvae hatched. 

Pupa (Fig. 18). 30 mm in length; shape like that of T. onolaus but color like that of 
T. catops. Duration, 18 days. 

The overall duration from the time that the eggs hatched to the emergence of the 
adults was 54 days. T. myops has been previously recorded in Papua New Guinea feeding 
on coconut (Cocos nucifera) and oil palm (Elaeis guineensis) both of the Palmae (Dept. 
Primary Industry, unpublished). 


DISCUSSION 


Cycads are known to be toxic and often lethal to cattle. Whiting 
(1963) discussed the toxicity of cycads in general, and Yang and Mick- 
elsen (1968) have shown that the husk of Cycas circinalis is toxic to 
rats. It is quite probable, therefore, that the larvae of T. onolaus, like 
many “pharmacophagous”’ butterflies (the Aristolochia-feeding swal- 
lowtails, for example), can sequester, and store, certain compounds 
(such as bitter alkaloids) which render them distasteful to birds and 
other predators. Their bright wine-red color suggests this. The larvae 


VOLUME 38, NUMBER 2 81 


of T. butleri, which also feeds on cycads, are also wine-red (T. Fenner, 
pers. comm.). 

It is possible that the larvae of T. catops and T. myops are more 
palatable to their predators, because their foodplants are not known to 
have toxic properties. Other species and their foodplants, which have 
not yet been mentioned but which are relevant to this discussion, in- 
clude Taenaris artemis Vollenhoven on coconut (Cocos nucifera: Pal- 
mae) and T. phorcas on tanget (Cordyline: Liliaceae) (T. Fenner, pers. 
comm.). Rosier (1960) has found the wine-red larvae of T. horsfieldii 
on Smilax (Smilacaceae) and, according to Corbet and Pendlebury 
(1978), the closely related genus Faunis in Malaysia feeds on Smilax 
(Smilacaceae), Musa (Musaceae) and Pandanus (Pandanaceae). Recent 
records of other Taenaris foodplants sent into the Insect Farming and 
Trading Agency include Taenaris dimona Hewitson on banana (Musa: 
Musaceae) and Taenaris gorgo Kirsch on Black Palm (Caryota rum- 
pha: Palmae). Both records were from the Maprik area, East Sepik 
Province. I have recorded the life history of Taenaris artemis on Pan- 
danus (Pandanaceae) in the Western Province. The larvae were pre- 
dominantly yellow marked with black. 

Although Taenaris larvae do not appear to advertise their presence, 
all species nevertheless feed gregariously, which is behavior character- 
istic of distasteful Lepidoptera. However, on some foodplants the lar- 
vae of certain Taenaris species may be unable to store adequate sec- 
ondary plant compounds for their effective protection. If T. catops 
obtains no such protection by feeding on ground orchids, then this may 
explain why adults were seen to imbibe cycad juices and consequently 
were so common in the study area. A similar conclusion was reached 
by Edgar et al. (1976) for danaine butterflies that enhanced their un- 
palatability by visiting the withered leaves of plants which produced 
pyrrolizidine alkaloids. The observation that T. onolaus, even as an 
adult, imbibed cycad juices strongly supports the hypothesis that Taen- 
aris is a distasteful group of butterflies and that some species enhance 
this as adults. It may be added that the fermenting skins of cycad nuts 
have an extremely nauseating smell. 

All species of Taenaris so far studied in the field have exhibited 
great wariness and are quick to avoid capture. This, together with their 
eyespots and the protective hairs and bristles of their larvae, may be 
considered to be secondary lines of defence if they have been retained 
from an ancestral form that was more cryptically colored and in which 
these characters were of primary protective function. Such an ancestor 
may have looked like the small, dull, species of Fawnis found in Ma- 
laysia today. The general trend to enhance the aposematic attributes 


82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of Taenaris appears to have been for the butterflies to increase in size, 
to become lighter, and for the eyespots to become enlarged and high- 
lighted with broader orange borders. Of the cycad feeding species so 
far studied, the ground color is predominantly black, and the extent 
of the orange has been greatly increased so that it is highlighted as a 
warning color. It is interesting to note that the same also appears to be 
true of the cycad feeding lycaenid Luthrodes cleotas from Papua New 
Guinea which has large patches of orange on the upper and underside 
of its hindwings in both sexes. 

It is possible that the ability of certain Taenaris species to feed on 
cycads as larvae is a recent evolutionary advance. J. Holloway (pers. 
comm. in discussion) suggested that the initial transfer to these prim- 
itive gymnosperms may have been a result of the similarity in the 
appearance of cycad fronds and those of coconut palms, for example, 
so that some Taenaris females began to oviposit on them by mistake. 
Alternatively, the transition from angiosperms, such as palms, to the 
cycad gymnosperms may have been through other angiosperms (such 
as Cordyline or Tapenochilus), that acted as “bridges,” i.e., they con- 
tained secondary substances that were common (or similar) to both. 
These may have acted either as oviposition cues to the females or 
phagostimulants to the larvae. The fact that adults of T. catops imbibe 
cycad juices could be taken to imply a closer link of this species with 
cycads in the past; however, it is also indicative of a chemical similarity 
between its normal foodplants and cycads. 

T. catops exhibits a wide range of geographical forms and it is 
probable that it is a Millerian mimic of its close relatives. A recent 
sampling of Taenaris in the Cape Rodney area, Central Province, 
revealed what appears to be a Miillerian mimicry complex involving 
four species of morphines (pers. obs., Dec. 1979). These were Hyantis 
hodeva Hewitson, Taenaris mailua Grose-Smith, T. catops and T. 
myops. They were all extremely alike, and in particular, T. catops was 
more heavily marked than usual with extended black margins to the 
apices of the fore and hind wings. T. mailua differed in the area from 
the form of the nominate race and was slightly less heavily marked 
with black. It appears, therefore, that there was a convergence of the 
phenotypes of all the species in the area. All four species looked iden- 
tical on the wing. 

Miillerian mimicry within the Morphinae appears to be a wide- 
spread phenomenon throughout New Guinea in general, and another 
good example has been recorded from Minj in the Western Highlands 
Province between H. hodeva and T. catops where the extremely white 
form of T. catops is predominant (Figs. 22 & 24). H. hodeva in the 
area is almost white, lacks its usual heavy black apical margins and has 


VOLUME 38, NUMBER 2 83 





Fics. 21-24. Miillerian mimicry in female morphines: 21, normal Taenaris catops 
from Bulolo; 22, albinotic T. catops from Minj; 23, normal Hyantis hodeva from Bulolo; 
24, albinotic H. hodeva from Minj. (Males from the two localities are like their females. ) 


reduced eyespots. In and around the Bulolo Valley the same species 
are also alike, but in this locality they are heavily marked (Figs. 21 & 
23) and conform to the more normal and widespread phenotypes. 
Other butterflies are probably Batesian mimics of Taenaris. For 
example, the female of Mycalesis drusillodes Oberthiir is thought to 
be mimetic of H. hodeva (Vane-Wright, 1971). Both model and mimic 
have been collected from the Torricelli Mountains near Maprik in the 
East Sepik Province (P. Clark, pers. comm.) and at Mt. Bosavi in the 
Southern Highlands Province (pers. obs., April 1980). The satyrine 
genus Elymnias is apparently mimetic of certain species of Euploea 
(Danainae), and Elymnias agondas Boisduval females are extremely 
good mimics of Taenaris bioculatus Guérin and T. catops where the 
models and mimics occur sympatrically. Hypolimnas deois Hewitson 
(Nymphalidae), in color and pattern, is very Taenaris-like and may 
be mimetic of T. onolaus in the Bulolo Valley. It is also assumed that 
the female form onesimus Hewitson of Papilio aegeus Donovan (Pa- 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pilionidae) mimics T. catops, and this form is also commonly seen 
around Bulolo. If this is so, then the female form amanga Boisduval 
of this swallowtail is a good candidate to be a mimic of T. onolaus. It 
has been seen frequently in the study area and strongly resembles T. 
onolaus in flight. 

A more detailed study of the foodplant relations and mimetic asso- 
ciations of these butterflies will prove most interesting as further life- 
histories and the foodplants of other species of Taenaris are discovered. 


ACKNOWLEDGMENTS 


I am most grateful to Ted Fenner (Department of Primary Industry, Darwin, Austra- 
lia), Dick Vane-Wright and Jeremy Holloway (British Museum of Natural History, Lon- 
don), all of whom have improved this manuscript by their most helpful suggestions. Also 
to Peter Clark (Insect Farming and Trading Agency, Bulolo, PNG) for his assistance in 
completing the rearing and the photography of T. myops. 


LITERATURE CITED 


BROOKS, C. J. 1950. A revision of the genus Tenaris Hiibner. Trans. R. Entomol. Soc. 
Lond. 101:179-238. j 

CorBET, A. S. & H. M. PENDLEBURY. 1978. The Butterflies of the Malay Peninsula. 
3rd edition revised by J. N. Eliot. Malayan Nature Society. 578 pp. 

D’ABRERA, B. L. 1977. Butterflies of the Australian Region. Landsdowne. 2nd edition, 
415 pp. 

EDGAR, J. A., P. A. COCKRUM & J. L. FRAHN. 1976. Pyrrolizidine alkaloids in Danaus 
plexippus L. and Danaus chrysippus L. Experientia 32:1535-1537. 

PyLe, R. M. & S. HUGHES. 1978. Conservation and utilization of the insect resources 
of Papua New Guinea. 157 pp. Unpublished mimeo. report, Wildlife Division, Papua 
New Guinea. 

Rosier, J. P. 1940. Aateekeningen over ontwikkelingsstadia van eenige javaansche 
vlinders. Entomol. Med. Ned.-Indie. 6:61-64. 

SIBATANI, A. & R. GRUND. 1978. A revision of the Theclinesthes onycha complex 
(Lepidoptera: Lycaenidae). Trans. Lepid. Soc. Jap. 29:1-34. 

SZENT-IVANY, J. J. H., J. S. WOMERSLEY & J. H. ARDLEY. 1956. Some insects of Cycas 
in New Guinea. P & NG. Ag. J. 11:1-4. 

VANE-WRIGHT, R. I. 1971. The systematics of Drusillopsis Oberthiir (Satyrinae) and 
the supposed Amathusiid Bigaena van Eecke (Lepidoptera: Nymphalidae), with 
some observations on Batesian mimicry. Trans. R. Entomol. Soc. Lond. 123:97-123. 

WHITING, M. G. 19638. Toxicity of cycads. Econ. Bot. 17:271-302. 

YANG, M. G. & O. MICKELSEN. 1968. Cycad husk from Guam: Its toxicity to rats. Econ. 
Bot. 22:149-154. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 85-87 


A NEW SPECIES OF SIMILIPEPSIS AND TAXONOMIC 
PLACEMENT OF THE GENUS (SESIIDAE) 


PING YUAN WANG! 


~ Research Entomologist, Institute of Zoology, 
Academia Sinica, Peking, China 


ABSTRACT. A new species of the wasp-like sesiid of the genus Similipepsis is de- 
scribed, and the taxonomic placement of this genus into the subfamily Tinthiinae is 
proposed. 


The Section of Entomology of the Carnegie Museum of Natural 
History (CMNH) maintains a large collection of insects that has been 
vastly underutilized by systematists. The collection is rich in all insect 
groups but butterflies and moths are particularly abundant. The di- 
versity of taxa is particularly evident among the collection of unsorted 
moths in which I found a sesiid specimen with remarkable ichneu- 
monoid resemblance. 

Further study of this wasp-like moth revealed that it belongs in the 
genus Similipepsis, a genus described by LeCerf (1911) and heretofore 
taxonomically unaligned in the Sesiidae hierarchy. Heppner and Duck- 
worth (1981:44) in their recent work made no study of this genus. 
They listed Similipepsis among other “‘unassigned” sesiid genera, leav- 
ing this problem for further research. 

My studies of the genus revealed that Similipepsis species are char- 
acterized by having the abdomen constricted to a slender pedicel at 
the base, the proboscis normal, labial palpus oblique with the second 
joint of long hairs, forewing veins R, and R; stalked and M, missing, 
hindwing with vein Cu, from just before angle of cell and widely 
separated from Cu,, hind leg wasp-like. The genus is further recogniz- 
able by the absence of the scale tuft on the tip of the antennae. Ac- 
cording to recent classification (Naumann, 1971; Duckworth & Eichlin, 
1977), these two characteristics suggest that Similipepsis has affinities 
and should be placed with genera of the subfamily Tinthiinae. 

To date, there are only four known species of Similipepsis, S. aurea 
Gaede, S. lasiocera Hampson, S. typica Strand and S. violaceus Le- 
Cerf. The genus is paleotropical in origin and is confined geographi- 
cally to the Ethiopian and Oriental regions. After reviewing specimens 
and literature of known species (Strand, 1913; Hampson, 1919; Gaede, 
1929), I determined that the aforementioned specimen in the Carnegie 


‘Resident Museum Specialist, Section of Entomology, Carnegie Museum of Natural History, USA. 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Adult male (holotype) of Similipepsis ekisi Wang, new species. 


collection collected from the Cameroons was quite distinctly different 
and not conspecific with the known species. 

I am indebted to Dr. Ginter Ekis for offering me the opportunity to 
study in the Section of Entomology. I would also like to express my 
appreciation to Dr. Chen Wen Young, the Collection Manager of the 
Section, for various courtesies during my six month research visit. I am 
indebted to Anna Tauber and Pat Vachino for literature and clerical 
assistance, and also to Vincent Abromitis, Section of Exhibits, for pho- 
tographic assistance. Dr. Craig Black, Director of the Carnegie Mu- 
seum of Natural History, provided the financial assistance that made 
it possible for me to come to the United States. 


Similipepsis ekisi, new species 


Holotype: Male Metet (Adamaoua), Cameroon (Republic of Cameroon), 15 August 
1919. A. I. Good. Carn. Mus. Acc. 6552 (deposited in CMNH, Holotype number 775). 

The holotype is associated with the following items: sex label (white, machine print); 
locality label (white, machine print); collection date label (white, machine and hand 
print); accession label (white, machine and hand print); CMNH repository label (yellow, 
machine print); holotype label (red, machine and hand print). 


VOLUME 38, NUMBER 2 87 


Male. Head: vertex black; frons brown; occipital fringe greyish white; vertex laterally 
with fringe greyish white mixed with black; labial palpi upturned, first and second 
segments brown and covered with extended long bushy scales, second segment less ex- 
panded than first, with brown scales on both sides and white erect scales on inside border, 
third segment white and sharply upturned above vertex; antenna brown and bipectinate, 
devoid of apical scale tuft; proboscis present. Thorax dark brown, tegula brown; meta- 
thorax with minute, slender brown and white hairy scales extended from base of hind- 
wing. Abdomen dark brown, first segment expanded slightly, second extremely narrowed 
and extended into a long stalk, third slightly expanded, fourth and fifth greatly expanded, 
sixth and seventh narrowed; anal tuft covered with setaceous, V-shaped brown scales; 
underside of third abdominal segment with ring of V-shaped white scales. Forewing 
transparent, except on costal margin; stem of R vein covered with dark brown scales, 
dark brown scales scattered in region of cell; cilia brown. Hindwing hyaline, with few 
scattered scales; veins and margins brown; cilia brown. Foreleg: front of femur, with 
long row of compressed brown scales; tibia brown with metallic sheen; tarsus with me- 
tallic blue setaceous scales on tarsomere; other tarsal segments white. Mesothoracic leg 
dark brown, scales metallic blue, green or red. Tarsus brown, with spiny scales on tar- 
somere, other tarsal segments with mixture of white and brown scales. Hindleg dark 
brown, with two pairs of long white spurs. Forewing expanse, 26 mm. Adult as shown 
in Fig. 1. 

Distribution: Known only from holotype from Metet (7°05'N, 13°17'E), Adamaoua, 
Cameroon, in Western Africa. 

Remarks: This species is superficially similar to S. violaceus. It differs from S. violaceus 
by the narrower costal margin. Also, the ventral side of the abdomen of the S. ekisi 
specimen with V-shaped white band which is distinctly absent in S. violaceus. 

This species is named in honor of Dr. Ginter Ekis, Curator of Section of Entomology, 
Carnegie Museum of Natural History at this writing. 


LITERATURE CITED 


DUCKWORTH, W. D. & T. D. EICHLIN. 1977. A classification of the Sesiidae of America 
North of Mexico (Lepidoptera: Sesioidea). Occas. Papers Entomol., Calif. Dept. Food 
& Agric. 26:1-54. 

GAEDE, M. 1929. Familie: Aegeriidae (Sesiidae) in A. Seitz, Die Gross-Schmetterlinge 
der Erde, II. Abteilung: Exotische Fauna, 14 (Die afrikanischen Spinner und 
Schwarmer):517-538. Plate 77. Stuttgart: A. Kernen. 

HAMPSON, G. F. 1919. A classification of the Aegeriidae of the Oriental and Ethiopian 
Regions. Novitates Zoologicae 26:46-119. 

HEPPNER, J. B. & W. D. DUCKworRTH. 1981. Classification of the superfamily Sesioidae 
(Lepidoptera: Ditrysia). Smithsonian Contrib. Zool., No. 314:1-144. 

LECERF, F. 1911. Descriptions d’Aegeriidae nouvelles. Bulletin du Museum National 
d Histoire Naturelle (Paris) 17:297-307. 

NAUMANN, C. M. 1971. Untersuchungen zur Systematik und Phylogenese der holark- 
tischen Sesiiden (Insecta, Lepidoptera). Bonner Zoologische Monographien (Bonn) 1: 
1-190. (English translation: 1977, Studies on the Systematics and Phylogeny of Hol- 
arctic Sesiidae (Insecta, Lepidoptera). 208 pp. Washington: Smithsonian Institution.) 

STRAND, E. 1918. Zoologische Ergebnisse der Expedition des Herrn G. Tessmann nach 
Siid-Kamerun und Spanisch-Guinea: Lepidoptera. IV. Archiv fiir Naturgeschichte 
(Berlin) 78A(12):30-84. 2 plates. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 88-91 


NOTES ON THE LARVA OF 
CARGIDA PYRRHA (NOTODONTIDAE) 


GEORGE L. GODFREY 


Illinois Natural History Survey, Natural Resources Building, 
607 E. Peabody, Champaign, Illinois 61820 


ABSTRACT. The ultimate instar larva of Cargida pyrrha (Druce) (Notodontidae) is 
described. Illustrations of its mandible and hypopharyngeal complex are included. The 
host plant of C. pyrrha is Condalia lycioides (A. Gray) Weberb.; an earlier report of 
Lycium may be in error. 


Comstock (1959) described the egg and several larval instars of Car- 
gida pyrrha (Druce), including what he called “... the third or fourth 
instar... .” (see his fig. 2). He succeeded in obtaining a subsequent 
larval instar, which he noted did not appreciably change in pattern or 
color from the preceding instar, but it is not certain that he was refer- 
ring to the ultimate instar. His larvae were reared from eggs that he 
found in Madera Canyon, Santa Rita Mountains, Arizona, on twigs of 
Lycium (Solanaceae), a questionable host identification. The eggs re- 
sembled those laid by captive Cargida pyrrha which he had observed 
earlier, and on that basis he identified the eggs and larvae subsequently 
described. However, no adults were reared to substantiate his judge- 
ment, and the identification cannot be definitely confirmed because of 
the apparent lack of preserved specimens. According to Donahue (pers. 
comm.), Comstock saved very few larval specimens during his work 
and did not give any larvae of C. pyrrha to the Los Angeles County 
Museum of Natural History, the main depository for his material. Based 
on the similarity of his figured “intermediate” instar from Madera 
Canyon to the mature larvae of C. pyrrha that the late R. G. Beard 
and I collected in 1967 (see below), it is assumed that his determination 
was correct. The purpose of this article is to comment on the host plant 
association of C. pyrrha and to describe the ultimate instar as a com- 
plement to Comstock’s earlier information on this species. 

Numerous mature larvae of Cargida pyrrha (Fig. 1) were seen de- 
foliating white crucillo, Condalia lycioides (A. Gray) Weberb. (Rham- 
naceae), on 31 July and 1 August 1967 at 4200 ft in Guadalupe Canyon 
(Cochise County) in the extreme southeastern corner of Arizona. Ac- 
tively feeding larvae were collected—several per bush at sunset on 
both evenings. The range of daily feeding activity is unknown. Within 
a few days they finished their larval feeding, and from 17 associated 
moths which emerged in 1968, J. G. Franclemont determined their 
identity. 


VOLUME 38, NUMBER 2 89 





Fic. 1. Cargida pyrrha, mature larva, Guadalupe Canyon, Cochise County, Arizona 
(photo by J. G. Franclemont). 


Comstock’s record of Lycium needs to be verified, because he may 
have misidentified the host. McFarland (pers. comm.) wrote that the 
plant illustrated by Comstock looks much like the figure of Condalia 
lycioides in Benson & Darrow (1954, fig. 60A) [Ziziphus obtusifolia 
(Hooker) A. Gray var. canescens (A. Gray) M. C. Johnston in Benson 
& Darrow (1981)] and like the C. lycioides growing in his own garden, 
partly because its leaves are not in fascicles. The leaves of Lycium are 
mostly fasciculate (Kearney & Peebles, 1960). McFarland added that 
the first time he saw Condalia lycioides he was sure that he was looking 
at a Lycium. 


Cargida pyrrha (Druce) 


General (Fig. 1): Total length 30-38 mm. Chaetotaxy basically noctuoid with extra 
setae only on lateral aspects of Ab8-6 prolegs. Head: smooth, width 3.3-3.8 mm. Body: 
integument smooth, velvety in appearance; dorsum of Ab8 slightly humped. All setae 
simple. Prolegs present on Ab3-6, 10; pairs on Ab3-6 subequal, Ab10’s slightly reduced; 
crochets in uniordinal, homoideous mesoseries. Midventral prothoracic glandular opening 
present. 

Hypopharyngeal complex (Fig. 2): Spinneret tapers distad, about twice length of Lps- 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





30 ——_ 


Fics. 2 & 8. Cargida pyrrha: 2, hypopharyngeal complex, left lateral view (Lp-1 = 
seta on first segment of labial palpus, Lp-2 = seta on second segment of labial palpus, 
Lps-1 = first segment of labial palpus, Lps-2 = second segment of labial palpus, S = stip- 
ular seta, Sp = spinneret); 3, left mandible, oral view (leader lines mark positions of the 
five outer teeth). Scale lines = 0.5 mm. Descriptive terminology follows Godfrey (1972). 


1, surpasses tip of Lp-2, dorsal margin of tip U-shaped in transverse cross section, distal 
lip entire. Stipular seta about half length of Lps-1, twice length of Lp-1 and also of Lps- 
2, subequal to Lp-2. Distal region separated from proximal region by shallow medial 
transverse cleft; spines absent proximate to spinneret, becoming numerous proximad. 
Proximolateral and proximomedial regions uniformly covered by short, thin spines. 

Mandible (Fig. 3): Outer teeth low, obtusely triangular, five teeth discernible (appear 
to be worn in available specimens); three large inner teeth present, flattened distally, 
adjoined by sinuate bridge. Two outer setae present (not visible in oral view of Fig. 3), 
insertions widely separated from each other. 

Coloration: Head black. Body ground color black; dorsal area black with two yellow 
stripes continuous to Ab8; subdorsal lines white; dorsal and ventral stripes of subdorsal 
area white and yellow respectively, both with black dorsal borders; lateral area yellow 
with black line above spiracles; ventral area mottled black and white with yellow line at 
level of prolegs and black midventral line; black patches above prolegs. Thoracic legs 
and prolegs orange. Spiracles black. 

Material examined: Six mature larvae: Guadalupe Canyon, 4200 ft, Cochise County, 
Arizona; 31 July & 1 August 1967; feeding on Condalia lycioides (A. Gray) Weberb.; G. 
L. Godfrey & R. G. Beard, collectors. Hypopharyngeal complex on GLG Slide 2500, 
John G. Franclemont Collection. 


The character of the three, large, adjoined inner teeth on the man- 
dible of Cargida pyrrha readily separates this species from other de- 
scribed notodontid larvae in the USA. Nothing comparable was found 
in the larvae of 26 other genera of North American notodontids that I 
recently examined. Note also that Gardner (1943) did not mention 
similar mandibular structuring in any of the Asiatic notodontid larvae, 
representing 14 genera, that he described. Perhaps this character will 
prove to be of some phylogenetic importance. However, notodontid 
larval mouthparts presently are too meagerly documented to evaluate 


VOLUME 38, NUMBER 2 91 


their phylogenetic implications, let alone draw conclusions from the 
mandible of a single species. 


ACKNOWLEDGMENTS 


I thank J. G. Franclemont, Cornell University, for his assistance and kind hospitality 
that made the completion of this paper possible. I also thank N. McFarland, Sierra Vista, 
Arizona, for his comments on Condalia and Lycium, and J. P. Donahue, Los Angeles 
County Museum of Natural History, for his information. P. A. Hyppio, Bailey Hortorium, 
Cornell University, provided the identification of Condalia lycioides. This project was 
funded in part by USDA AGR RMA Grant No. 12-14-100-8031-(33), Franclemont, Prin- 
cipal Investigator, and by support from the Illinois Natural History Survey. Preliminary 
drafts of the manuscript were reviewed by Franclemont, McFarland, L. M. Page, J. D. 
Unzicker, and D. W. Webb. 


LITERATURE CITED 


BENSON, L. & R. A. DARROW. 1954. The Trees and Shrubs of the Southwestern Deserts. 
2nd ed. The University of Arizona Press, Tucson and The University of New Mexico 
Press, Albuquerque. 437 + x pp. 

1981. Trees and Shrubs of the Southwestern Deserts. 3rd ed. The University 
of Arizona Press, Tucson. 416 + xviii pp. 

ComsTOCcK, J. A. 1959. Rare or common! With notes on the life histories of two south- 
western moths. Bull. South. Calif. Acad. Sci. 58:155-161. 

GARDNER, J.C. M. 1943. Immature stages of Indian Lepidoptera (5). Indian J. Entomol. 
5:89-102. 

GopFREY, G. L. 1972. A review and reclassification of larvae of the subfamily Had- 
eninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S.D.A. Tech. Bull. 
1450, 265 pp. 

KEARNEY, T. H. & R. H. PEEBLES. 1960. Arizona Flora. University of California Press, 
Berkeley and Los Angeles. 1085 + viii pp. 





Journal of the Lepidopterists’ Society 
38(2), 1984, 92-95 


THE LARVA OF AUTOGRAPHA FLAGELLUM (WALKER) 
(NOCTUIDAE: PLUSIINAE) 


KENNETH NEIL 


Department of Biological Sciences, 
Simon Fraser University, Burnaby, B.C. V5A 1S6 


ABSTRACT. The mature larva of Autographa flagellum (Walker) is described and 
illustrated. 


The noctuid genus Autographa Hiibner (Plusiinae) is represented in 
North America by sixteen species (Eichlin & Cunningham, 1968). Lar- 
val descriptions of only eight species have been published to date. The 
larvae of our known species of Autographa are all semi-loopers, lacking 
prolegs on abdominal segments 3 and 4. In the past, identification of 
all our nearctic Plusiinae has been difficult due to the small number 
of reliable differentiating characters available. Recent investigations 
have shown that the larval mouthparts, especially the hypopharyngeal 
complex, offer good separating characters (Eichlin & Cunningham, 
1969, 1978). The larvae have been reported as feeding on a wide 
variety of low plants and trees (Eichlin & Cunningham, 1978). 

Autographa flagellum was described by Sir Francis Walker in 1857 
from material collected at St. Martins Falls, Ontario. A. flagellum is a 
boreal species distributed from Newfoundland west to Alberta (Forbes, 
1954) and British Columbia (Llewellyn-Jones, 1951) and in the east, 
south to Maine and New Hampshire (Eichlin & Cunningham, 1978). 
Although Tietz (1972) lists Helianthus sp. and Liatris sp. as host plants, 
no description of the immature stages has been published. 

Ova were obtained from a female A. flagellum collected on 4 July 
1978 at Belliveau Cove, Digby County, Nova Scotia. Larvae were fed 
an artificial diet based on that of Hinks and Byers (1976). All larvae 
grew quickly and had pupated by 1 September 1978. Adults emerged 
1-10 October. Under natural conditions, A. flagellum overwinters as 
a third or fourth instar larva, as do most northern Plusiinae (Eichlin & 
Cunningham, 1978). 

This paper describes the mature larvae of Autographa flagellum. 
The terminology and abbreviations used follow Godfrey (1972) and 
Eichlin and Cunningham (1969, 1978). All illustrations were drawn to 
scale using a camera lucida and stereomicroscope. 


Autographa flagellum (Walker) 


General. Head 2.4—2.9 mm wide. Total length 33.5-36.1 mm. Head and body smooth, 
no microspines or granules present. No vestige of prolegs present on Ab3-4. Prolegs on 


VOLUME 38, NUMBER 2 93 





Fics. 1 & 2. Autographa flagellum: 1, dorsal view; 2, lateral view (x3). 


Abd same size as those on Ab6. Crochets biordinal, 21-23 per fifth and sixth abdominal 
proleg. All setae simple. 

Coloration (living material). Head (Fig. 3): Yellowish brown, no coronal freckles or 
reticulations present. Body (Figs. 1 & 2): Green with white flecks, flecks heaviest in 
subdorsal, lateral, and ventral areas; ventral half of subdorsal area on T1-8 darker green; 
dorsal margin of lateral area white; middorsal line green with narrow white line on 
edges; white lines obsolete on T1; subdorsal area green with narrow white line on dorsal 
and ventral edges; lines on dorsal edge obsolete on T1-3. Spiracles yellowish brown with 
darker brown peritremes. Lateral shield of prolegs green, becoming yellowish brown 
distally. Thoracic legs yellowish brown, darker brown distally. Prothoracic shield yellow- 
ish brown. 

Head (Fig. 3). Epicranial suture 0.59-0.64 mm long; height of frons (apex to Fa’s) 
0.75-0.78 mm; distance from F1 to anterior edge of clypeus 0.40—0.42 mm; interspace 
between F1—-F1 0.31-0.34 mm; AFa anterior and AF2 posterior to apex of frons; Al—A3 
forming an obtuse angle at A2; P1—P1 0.67-0.71 mm, P2-P2 0.85-0.87 mm. Distance 
from Pl to epicranial suture about % that from Pl to L; L posterior of juncture of 
adfrontal ecdysial line. Ocellar spacing: Ocl-—Oc2 0.07-0.09 mm; Oc2-—Oc3 0.05-0.07 
mm; Oc3-—Oc4 0.038-0.04 mm. 

Mouthparts. Hypopharyngeal complex (Fig. 4): Spinneret elongate, tapering distally, 
subequal to Lp2; Lpsl shorter than Lp2; Lps2 less than % the length of Lpl; stipular 
setae extremely short, about /, length of Lpsl, 4, length of Lpl, and about \, length of 
Lp2; distal and proximal regions of hypopharyngeal complex continuous; distal 4 bare, 
remainder spined, spines becoming much longer and more robust proximally. Raduloid 
with 18 ridges. Mandible (Figs. 5 & 6): Two well-separated outer setae present; inner 
surface with three distinct ridges and small, acutely angled inner tooth on rib 2; six outer 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


EM yp —— 

















Fics. 3-8. Autographa flagellum: 3, head capsule, frontal view; 4, hypopharyngeal 
complex, left lateral view; 5, left mandible, oral surface; 6, left mandible, outer surface; 
7, anal shield, dorsal view; 8, dorsolateral chaetotaxy of prothoracic (T1), mesothoracic 
(T2), and abdominal segments (Ab1-2 and Ab6-10). Scale lines = 1 mm. 


teeth present, first small, second to fifth well-developed and angular; second to fourth 
serrated on both sides; sixth outer tooth about twice width of fifth, low and rounded. 
Thorax. Segment T1 (Fig. 8): Prothoracic shield smooth and weakly sclerotized; SD1 
and SD2 setal insertations well-separated from shield; SD1 very fine and hairlike with 
thickened sclerotized annulus at base; L2 as in SD1 but with annulus thinner and less 
heavily sclerotized; interspace D1-D1 about 0.67 XD1-XD1; D1-SD2 about 1.34 SD2- 
X D2; spiracle transversely aligned with posterior margin of prothoracic shield, elliptical, 


VOLUME 38, NUMBER 2 95 


0.21-0.23 mm high, 0.13-0.14 mm wide; peritreme wider laterally; coxal bases contig- 
uous. Segments T2-3 (Fig. 8): SD1 very fine, but with annulus narrower and less heavily 
sclerotized than on T1; SV2 obsolete; all setae hairlike, tapering and sharply pointed; 
coxal bases narrowly separated. 

Abdomen. No vestige of prolegs present on Ab3—4. Dorsal and lateral chaetotaxy of 
Ab1-10 as in Fig. 8. SD1 thicker, not fine as on T1-3. Ab] and Ab7-8 with SV2 lacking, 
Ab2-4 with SV1 and SV2 partially fused. Ab4: V1-V1 close together; V1-SV2 about 
twice the distance of V1-V1. Ab10: Anal shield as in Fig. 7. Dorsal margin convex, 
posterior margin entire. Length of D1 on Ab6-7 0.57-0.61 mm; D2 0.71-0.72 mm. Asp7 
0.22-0.25 mm high, 0.13-0.14 mm wide; Asp8 0.35—-0.36 mm high, 0.17-0.20 wide. 

Material examined. 10 specimens: Belliveau Cove, Digby County, Nova Scotia. Reared 
on artificial diet (Hinks and Byers, 1976) from ova obtained from a female collected on 
4 July 1978. Adults emerged 1-10 October 1978. Moth collected, determined, and larvae 
reared by K. A. Neil. 


Remarks 


Based on the key to the known species of Plusiinae larvae given by 
Eichlin and Cunningham (1978), A. flagellum appears closest to A. 
rubida Ottolengui, as both species have a tooth on the second mandib- 
ular ridge. A. flagellum differs from that species, however, by its over- 
all green color including the head, the smooth integument, the V1-V1 
interspace on Ab4, and by the raduloid which has 18 ridges. 


ACKNOWLEDGMENTS 


I thank Dr. T. D. Eichlin of the Insect Taxonomy Laboratory, Sacramento, California 
for reviewing this manuscript and Ronald Long of Simon Fraser University for the 
photography. 


LITERATURE CITED 


EICHLIN, T. D. & H. B. CUNNINGHAM. 1969. Characters for the identification of some 
common Plusiinae caterpillars of the southeastern United States. Ann. Entomol. Soc. 
Amer. 62(3):507-510. 

1978. The Plusiinae (Lepidoptera: Noctuidae) of America north of Mexico, 
emphasizing genitalic and larval morphology. U.S. Dept. Agr. Tech. Bull. 1567: 
122 pp. 

ForBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Pt. III. 
Cornell Univ. Agr. Expt. Sta. Mem. 329:433 pp. 

GopFreEy, G. L. 1972. A review and reclassification of larvae of the subfamily Had- 
eninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agr. Tech. 
Bull. 1450:265 pp. 

HINKS, C. F. & J. R. Byers. 1976. Biosystematics of the genus Euxoa (Lepidoptera: 
Noctuidae). V. Rearing procedures and life cycles of 36 species. Can. Entomol. 108: 
1345-1357. 

LLEWELLYN-JONES, J. R. J. 1951. An annotated check list of the Macrolepidoptera of 
British Columbia. Entomol. Soc. B.C. Occasional Paper 1:148 pp. 

TiETZ, H. M. 1972. An Index to the Described Life Histories, Early Stages, and Hosts 
of the Macrolepidoptera of the Continental United States and Canada. 2 vols. A. C. 
Allyn, Sarasota, Florida. 1041 pp. 





Journal of the Lepidopterists’ Society 
38(2), 1984, 96-101 


A NEW HAWKMOTH FROM QUINTANA ROO, MEXICO 


VERNON ANTOINE BROU, JR.’ 
Rt. 1, Box 74, Edgard, Louisiana 70049 USA 


ABSTRACT. A new species of Manduca Hubner is described in its adult stage. This 
species is similar in maculation to Manduca morelia (Druce) and M. pellenia (Herrich- 
Schaffer). Differences in size, wing shape and genitalia prove it to be distinct. 


Manduca wellingi, new species 


(Figs. 1-5) 


Wing length. Males: 41 mm (35-44 mm, n = 64); females: 46 mm (44-50 mm, n= 
10). M. wellingi two-thirds size of morelia (Druce). Rothschild and Jordan (1903:79) 
treated morelia as being synonymous with pellenia (Herrich-Schaffer). A review of the 
original description of morelia makes it clear that this is incorrect. 

Wing maculation and shape. Color and maculation of wellingi very similar to those 
of morelia. Both species show about same degree of color variation when series compared. 
Forewing of wellingi light, sandy brown to tawny, with confluent series of black lustrous 
patches in median space, forming large, semicircular band, ends of which intersect costal 


1 Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer 
Services. 





Fic. 1. Manduca wellingi, n. sp. Holotype, male, Nuevo X-can, Quintana Roo, Méx- 
ico. 27 Sept. 1981 (E. C. Welling M.; U.S. N.M.N.H.). 


VOLUME 38, NUMBER 2 97 


Fic. 2. Wing outline comparison of males (a) Manduca wellingi, (b) Manduca mo- 
relia, (ec) Manduca pellenia. 


margin. Most evident are crescent shaped, off-white stigma, distinct, oblique, zig-zag, 
black apical dash, crenulate subterminal line that essentially parallels postmedial band, 
which, as in morelia, consists of two, closely parallel lines. Both postmedial and ante- 
medial bands double and may appear triple as they are paralleled on side toward median 
space by usually less distinct, diffuse, mesial lines. Base pale, with one or two minute 
tufts of nearly white scales, enclosed by dark basal band that may be bisected by short, 
diffuse basal dash. Basal space with dark spot near middle. Nearly black hindwing crossed 
by two pale bands corresponding in position to antemedial and postmedial. Outer one 
runs from anal angle to costal margin just beyond middle, and inner one is small curved 
band enclosing black basal area. Light-brown outer margin becomes obscured distally. 
Forewing beneath is dark gray. In males, entire dark ventral surface heavily sprinkled 
with light-brown scales except for outer marginal band where light colored scales are 
wanting. Hindwing below exhibits same features, although light-brown scales dominate 
entire surface to much greater degree. Dark outer marginal band present and more 
distinct than on forewing. In females, amount of light-brown scales greatly reduced or 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 38. Genitalia, Manduca wellingi, male. 


nearly absent on underside of forewing, and outer marginal band is hardly distinguish- 
able. Hindwing similar to that of males. Both sexes have set of three sinuous dark median 
lines which arise at common point at anal angle of hindwing and transgress costal margin 
of forewing. Wing shape of wellingi clearly different from that of pellenia and morelia 
(Fig. 2). 

Other maculation. Thorax dorsally off-white, ventrally dark brown, completely sur- 
rounded by heavy black line. Tegula and head essentially concolorous with forewings. 
Palpus black on segment 8, but segments 2 and 1 mixture of light and dark brown. 

Abdomen typical for genus, with dorsal area similar in color to forewings, with distinct, 
full-length, medial black line; laterally black with large yellow patch on segments. Ob- 
solescent sixth spot, more noticeable in females, sometimes evident. Below and between 
each yellow patch are small, narrow, white intersegmental bands. Ventral surface off- 
white with brown scales throughout and usually with medial row of 1 to 4 small black 
spots. 

Genitalia. Genitalia of pellenia have been illustrated in Rothschild and Jordan (1903) 
and Mooser (1940) and are sufficiently different not to be confused with those of either 
morelia or wellingi. 

In wellingi (Fig. 3) apex of sacculus serrated and sometimes narrower than in morelia. 
Process of sacculus acuminate and curved inward more in wellingi. Hooked apex of 
gnathos minimal, unlike that of morelia, which has pronounced hooked apex. In female 
of wellingi (Fig. 4), lamella postvaginalis large and posterior margin only slightly in- 
dented. Lamella postvaginalis of morelia (Fig. 5) reduced in size and posterior margin 
strongly emarginate, unlike wellingi. 

Flight period. Adult specimens have been taken each month from 2 April to 2 No- 
vember, the greatest number being recorded during June. 

Types. HOLOTYPE 6 (Fig. 1) Nuevo X-cA4n, Quintana Roo, MEXICO. 27 Sept. 1981, 
E. C. Welling M. collector. USNM type no. 100721. ALLOTYPE 2, same locality, 27 July 


VOLUME 38, NUMBER 2 99 





wellene i 


Fic. 4. Genitalia, Manduca wellingi, female. 


1979, E. C. Welling M. PARATYPES: Same locality, 1 6, 1 2, 2 Sept. 1960, 1 4, 2 July 1963, 
1 6, 1 2, 15 Aug. 1963, 1 4, 6 June 1967, 1 6, 9 June 1967, 1 4, 29 July 1970, 1 4, 12 June 
1971; 1 3, 14 June 1971, 1 6, 25 Sept. 1975, 1 6, 10 June 1974, 1 46, 12 Sept. 1974, 1 6, 5 
June 1975, 1 6, 1 Nov. 1976, 1 4, 15 June 1977, 1 4, 15 July 1977, 1 6, 24 Aug. 1977, 1 
6, 15 July 1979, 1 4, 2 Aug. 1979, 2 64, 5 June 1980, 1 6, 10 June 1980, 1 4, 18 June 1980, 
1 6, 20 June 1980, 1 4, 9 July 1980, 1 4, 1 Aug. 1980, 1 4, 2 Aug. 1980, 1 6, 15 Aug. 1980, 
1 6, 28 Aug. 1980, 1 6, 29 Aug. 1980, 3 66, 1 Sept. 1980, 1 6, 2 Oct. 1980, 1 4, 1 Apr. 
19815 1 9, 5 Apr. 1981, 1 6, 25-Apr. 1981, 1 6, 27 Apr. 1981, 1 9, 4 June 198]; 1 6, 9 June 
198i, 1 6, 12 June 1981, 3 66, 18 June 1981, 1 6, 1 9, 22 June 1981, 1 6, 1 2, 25 June 
1981, 1 4, 26 June 1981, 1 4, 1 2, 28 June 1981, 2 44, 29 June 1981, 1 2, 28 June 1981, 1 
6, 29 June 1981, 1 4, 1 2, 1 July 1981, 1 4, 3 July 1981, 1 4, 7 July 1981, 1 4, 9 July 1981, 
1 4, 20 July 1981, 1 4, 30 July 1981, 1 4, 19 Aug. 1981, 1 6, 13 Sept. 1981, 2 68, 17 Sept. 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





norek/a 


Fic. 5. Genitalia, Manduca morelia, female. 


1981, 1 8, 20 Sept. 1981, 1 2, 22 Sept. 1981, 1 4, 1 Oct. 1981. Tintal, Quintana Roo, 
México, 2 64, Sept. 1976. Chetumal, Quintana Roo, México, 1 6, 11 May 1977, 1 6, 12 
May 1977. Tikal, E] Petén, GUATEMALA, 1 4, 18 July 1981. 

Holotype and allotype deposited in the U.S. National Museum of Natural History; 
paratypes in American Museum of Natural History, British Museum of Natural History, 
Universidad Central de Venezuela, Instituto de Biologia, México, D.F., México, and in 
the collections of E. C. Welling M. and V. A. Brou. 


ACKNOWLEDGMENTS 


I wish to thank Dr. D. C. Ferguson, Systematic Entomology Laboratory, USDA, Wash- 
ington, D.C. for manuscript review and helpful suggestions. Special thanks go to Sr. E. 


VOLUME 38, NUMBER 2 101 


C. Welling M., Mérida, Yucatan, México for supplying the entire series of specimens of 
this new species. 


LITERATURE CITED 


Druce, H. 1894. Ann. Mag. Nat. Hist., series 6, 13:169. 

GEHLEN, B. 1931. Ent. Zeitschr. Frankfurt a. M. 41:201. 

HERRICH-SCHAFFER, G. A. W. 1864. -Ausl. Schm. p. 50. 

MooseER, O. 1940. Anales Escuela nacional ciencias biologicas (Mexico) 1:407—495. 

ROTHSCHILD, W. & KARL JORDAN. 1903. A revision of the lepidopterous family Sphin- 
gidae. Novitates Zoologicae 9(supplement). 


Journal of the Lepidopterists’ Society 
38(2), 1984, 102-113 


NATURAL HISTORY NOTES FOR TAYGETIS ANDROMEDA 
(CRAMER) (SATYRIDAE) IN EASTERN COSTA RICA 


ALLEN M. YOUNG 


Invertebrate Zoology Section, Milwaukee Public Museum, 
Milwaukee, Wisconsin 53233 


ABSTRACT. The early stages and a larval food plant are reported for the first time 
for Taygetis andromeda (Cramer), as studied in eastern Costa Rica. Various aspects of 
larval and adult behavior, including feeding and egg-placement, are also discussed for 
this widespread Central American satyrid. The larval food plant is a grass, Acroceras 
zizanioides (Graminae). 


Taygetis andromeda (Cramer) is a common butterfly (Fig. 1) of 
forest and old secondary habitats from 0 to 1500 m on the Pacific and 
Caribbean watersheds of Costa Rica (A. M. Young, pers. obs.; P. J. 
DeVries, pers. comm.). The butterfly is also widespread throughout 
much of Central America and South America. It is one of four species 
of Taygetis reported from Central America, along with different species 
in Mexico, Central America, and South America which often exhibit 
distinctive patterns of range separation by habitat and elevation (Ross, 
1976; Ebert, 1969; Lamas, 1967; P. J. DeVries, pers. comm.). While 
some fragmentary information on the early stages of South American 
Taygetis other than andromeda exist (Muller, 1886; d’ Almeida, 1922), 
the present paper constitutes the first report of early stages and larval 
food plant for T. andromeda. 


METHODS 


The majority of observations were made at two localities on the 
Caribbean or Atlantic watershed of eastern Costa Rica: “Finca La 
Tigra,’ near La Virgen (10°23’N, 84°07’ W; 220 m elev.), Heredia Prov- 
ince (described as “premontane tropical wet forest’’); “Finca Experi- 
mental La Lola,” near Siquirres (10°06’N, 83°30’ W;; 30 m elev.), Limon 
Province (described as “lowland tropical wet forest”). At “La Tigra” I 
observed T. andromeda adults on a pile of rotting bananas placed 
along a footpath through old secondary forest adjacent to a cacao plan- 
tation; the bait was used at wide intervals between 1977 and 1982 to 
observe this species and other butterflies. At ““La Lola’ I witnessed 
egg-placement (oviposition) behavior in T. andromeda and conducted 
a study of the early stages by confining recently deposited eggs in large 
clear plastic bags along with fresh cuttings of the larval food plant. 
These cuttings were replaced every two or three days, and the bag was 
kept tightly shut. A voucher specimen of the food plant was collected 


VOLUME 88, NUMBER 2 103 





Fic. 1. Taygetis andromeda, reared from eastern Costa Rica: dorsal (left photo) and 
ventral (right photo) aspects of the adult. 


for determination. A careful check of early stages being reared was 
made to describe each stage, including estimates in days of duration 
for each stage. The bulk of the rearing was conducted at La Lola, with 
conditions in the rearing bag being of room temperature, which was 
the same as the air temperature in the nearby “La Lola’ cacao plan- 
tation. 


RESULTS 
Adult Natural History 


Taygetis andromeda adults readily come to rotting bananas on the 
ground in wet forest. Throughout the year at “La Tigra,” there is 
usually a mix of “worn” and “fresh” butterflies, suggesting a contin- 
uously breeding population here. The butterfly is often seen on bait 
with several other butterflies, including Morpho peleides limpida But- 
ler, M. granadensis polybaptus Butler (see also Young, 1982 for data 
on the abundances of these two species on banana baits at “La Tigra’’) 
(Morphidae), Caligo memmon Cramer, Caligo atreus uranus H.-Schaff. 
(Brassolidae), Caerois sp. (Satyridae), Nessaea aglaura Feld., Myscelia 
ethusa Bsdv., and Prepona spp. (Nymphalidae). Within the shaded 
forest understory, T. andromeda, in my experience, generally flies 
within one meter of the ground. Males are far more common at baits 
than females, a condition reflected also in the fact that males mostly 


104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


collected at baits are more commonly represented in museum collec- 
tions for this species. 


Egg-Placement Behavior and Larval Food Plant 


On 22 July 1982 at 1400 h, I observed T. andromeda ovipositing in 
an approximately 100 m? patch of grasses (canopy height 0.5-1.0 m) 
in a “light gap” within the “La Lola” cacao plantation. The butterfly 
alighted cross-wise on a broad blade of the bamboo-like grass used as 
a larval food plant here. The butterfly then curled the abdomen around 
to the underside and placed a single egg on the blade. She then flew 
off and repeated the behavior on nearby individuals of the same plant. 
The larval food plant, which is also the egg-placement site, is the grass 
Acroceras zizanioides (H.B.K.) Dandy (Graminae), as determined from 
fresh specimens by Dr. Richard Pohl. This grass species is common in 
the lowland and premontane wet forest regions of Costa Rica (R. Pohl, 
pers. comm.). An egg is seldom affixed to the same general area of a 
grass blade both in the field and when a butterfly is confined to a 
plastic bag with fresh cuttings of A. zizanioides (Fig. 2). Under con- 
fined conditions, several eggs are sometimes placed on a single grass 
blade (Fig. 2), and I obtained a total of 15 eggs in four days by this 
method. Many eggs are also scattered singly on grass blades under this 
condition. In the field, T. andromeda is active in the late afternoon 
and at dusk; possibly, the butterfly is also nocturnal, but this behavior 
has not been studied. During other hours of the day, the butterfly is 
readily “flushed out’? from palmaceous undergrowth in wet forest. 


Early Stages 


Egg. The egg is spherical, about 1.3 mm in diameter and very pale (almost white) 
green (Fig. 2). With a 10x hand lens, a very fine surface sculpturing, somewhat resem- 
bling the “hexagonal” pattern reported for the egg of T. ypthima by Muller (1886), is 
barely visible. Within a day or two of hatching, the black head capsule of the larva is 
clearly visible (Fig. 2), and the rest of the egg assumes the pale green color of the larva. 

First instar larva. The first instar is about 7 mm long just after hatching, and the body 
is cylindrical and straight, with a gradual tapering towards the posterior end (Fig. 3). 
The head capsule is shiny black and conspicuously “lobed” laterally making it wider 
than the trunk (Fig. 3). The head capsule has six prominent horn-like protuberances 
arising laterally from the lobed areas (Fig. 4). These are structures that disappear in later 
instars (Fig. 4). The frontal area of the head capsule has some small black setae, as also 
noted by Muller (1886) for T. ypthima, and a conspicuous patchwork of irregular “fis- 
sures’ to either side of the epicranial suture. The trunk is pale green and covered with 
almost translucent fine setae. The forked tail points upward at an angle of about 35°; 
each fork shaft is pink with a black tip. The terminal trunk segment and anal plate are 
white. 

Second instar larva. Just after the molt from the first instar, the larva is about 10 mm 
long. The trunk region is now uniformly light green with several longitudinal yellow 
stripes. The head capsule is strikingly different from that of the previous instar: it is bi- 
lobed in general appearance, and light green with yellowish stripes (Fig. 3). The larva 


VOLUME 38, NUMBER 2 


Fic. 2. Egg stage of Taygetis andromeda, and first instar larva feeding; clockwise, 
from upper left photo: three eggs scattered on a single grass blade of the larval food 
plant, Acroceras zizanioides (Graminae) as the result of female butterfly ovipositing in 
captivity; close-up view of one egg; egg position by female ovipositing in the field, and 
first instar larva feeding at the edge of a grass blade; first instar larva in initial stages of 
hatching. 





106 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


reaches a length of about 16 mm in three days (n = 6 larvae measured), whereas the first 
instar larva grows to 9 mm in the same amount of time. 

Third instar larva. The general appearance of the third instar larva is very similar to 
that of the previous instar, but the body assumes a thicker profile, and the annulets on 
the body segments are more prominent. The strongly bi-lobed head capsule (Fig. 4) is 
now pink, with the protuberances being sculptured and adorned with setae. The basic 
color pattern and head capsule structure (Fig. 4) of this instar is retained in the subse- 
quent two instars. The third instar larva grows to about 28 mm in three days (n= 4 
larvae measured). 

Fourth instar larva. This instar (Fig. 5) is very similar to the previous one, and it 
attains a body length of about 32 mm in five days (n = 4 measured). The head capsule 
is directed more anteriorly now and bears a pair of very prominent protuberances or 
tubercles (Fig. 5). The trunk is very noticeably arched (Fig. 5). 

Fifth instar larva. This instar is similar to previous ones (Fig. 6) but with considerable 
change in the configuration of the ornate head capsule (Fig. 4). As in the previous two 
instars, the background color of the trunk region is bright green with brown latero- 
ventral (supraspiracular) longitudinal stripes. The head capsule is tan frontally and with 
a dark brown thick vertical stripe on each side (lateral area). The tan color of the head 
capsule in the fifth instar larva replaces pink in the previous two instars. The posterior 
edges of the two prominent “horns” (Fig. 6) are tan, while dark brown both laterally 
and frontally. The dark brown areas of the “horns” extend down the sides of the head 
capsule. The broad, latero-ventral stripe on the trunk begins at the first body segment, 
being at first very faint on the first two segments, and then becoming much darker on 
later segments. This “stripe” is really a composite of two thick, dark brown lines sur- 
rounding a thin, central tan or cream-colored line, the latter barely visible, even with a 
10x hand lens. Dorsally the trunk bears another complex pattern of longitudinal stripes: 
thick lines of green alternate with faint streaks of pink, and the forked tail is also green, 
and about 5 mm long. The pinkish red longitudinal bands run dorsally; each band is 
tapered, about 20 mm long and 2 mm wide at the thickest point, and extends from the 
third to terminal abdominal segments. On the fourth segment, the band on each side is 
adorned with small, irregularly-shaped black markings, each composed of two parts: a 
larger oval area anteriorly, followed by a smaller one. Dorsally the thoracic area bears a 
transverse “ring” of irregular, black markings at the posterior edge of the third segment. 
This band blends into a few thin longitudinal black lines extending from the third 
segment anteriorly to the head capsule. The tapered profile, segmental annulets, and 
overall arched trunk region make the fifth instar very easy to recognize, along with the 
description of stripes (Fig. 6). The trunk region is covered with a fine, light brown or 
tan down of setae. The fifth instar grows to about 55 mm in 12 days (n = 4 measured). 

Pupa. The larva assumes a “J”’ position, undergoing little change in color, but with a 
contraction of body length to about 30 mm, and then molts within a day to the pupa 
stage (Fig. 7). The pupa is leaf-green all over and very stout in profile; it appears to be 
“dusted” with a waxy, whitish coat, more evident in some areas than in others. The pupa 
is 21 mm long by 9 mm thick (dorso-ventral axis through the thoracic area) and 9 mm 
wide (laterally, also through the thoracic area). A pale fulvous ridge defines the rear 
marginal areas of the forewing, and there is a pale, whitish blue thin line just below the 
spiracle area. The spiracles are marked with black. The abdominal area has dorsal, faint, 
multiple, longitudinal streaks of light green alternating with dark green lines. Of these, 
the medial, light green line is the thickest. There is a pair of pale yellow dots marking 
the beginning of two lateral thick whitish lines on the abdomen. These dots are on the 
first abdominal segment. The lateral and dorsal areas of the thorax have doublets of 
small, raised, pale yellow, dots; similar dots also occur immediately adjacent to the wing 
pads. The thoracic area bears a prominent longitudinal ridge which is pale fulvous at 
the apex. The dorsal area of the head capsule is slightly bi-lobed in the transverse plane. 
About 4% down the leg-case area from the head, there are two lateral pairs of irregularly- 
shaped white blotches followed by a pair of raised black dots. The wing pads also have 
several raised dots: one fulvous dot in the subcostal cell of the forewing; one black dot 
at the distal end of this cell; a whitish blotch between two radials; a small white dot on 


VOLUME 38, NUMBER 2 


: 
. 
‘ 
: 
= 
’ 
> 
I 
. 
4 
© 
i 
hf 
o> 


Fic. 3. First and second-instar larvae of T. andromeda, emphasizing overall body 
profile and some details of the head capsule. Clockwise, beginning with upper left photo: 
lateral view of first-instar larva; head capsule of first instar; frontal view of head capsule 
in the second instar larva; dorsal view of second instar larva. 





108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1mm 1mm Imm 


Fic. 4. Schematic drawings of head capsules, all frontal aspects, for three larval 
instars of T. andromeda. From right to left: first, third, and fifth instars, respectively. 
All three drawings made with drawing-tube attachment to Wild MS binocular micro- 
scope. 


the subcostal vein near the apex of the wing. The cremaster is green with a brown “hook” 
for attachment. For three pupae, all adults eclosed (Fig. 7) by 0800 h. On the afternoon 
and evening prior to eclosions, the pupa darkens considerably, this process beginning in 
the wing pads. Eclosion is very rapid, with the butterfly fully expanding its wings within 
ten minutes after leaving the pupal shell. 


Egg and Larval Natural History 


The life cycle requires about 48 days from egg to adult, with the 
egg lasting seven days, the larva about 26 days, and the pupa 15 days. 
As with many other Neotropical butterflies in which individual eggs 
are scattered among many food plant individuals in an area of habitat, 
the first instar larva of T. andromeda devours its emptied egg shell 
down to the base immediately upon hatching. In those Lepidoptera 
which cluster eggs on the food plant, such behavior is conspicuously 
absent, a trait that appears to function to prevent cannibalism of late- 
hatching eggs by the larvae of early-hatching eggs in the same cluster. 
After devouring the egg shell, the larva moves to the edge of the same 
grass blade and begins feeding on plant tissue and does so from the 
ventral surface of the blade (Fig. 3). All instars, but most noticeable in 
the first instar, have the habit of “shooting out” fecal pellets to a 
distance of about 1-5 cm from the feeding site. When eating, larvae 
of all instars perch on a thin silk matting on the food plant. The bulk 
of feeding in all instars is nocturnal, and individual larvae construct 
silken strands to and from feeding sites. Older instars (instars IV—V) 
often rest on grass stems rather than on blades when not feeding. The 
long, slender body profile of the earlier instars (I-III) give the larvae 
a cryptic appearance on grass blades; the arched appearance so evident 
in the older instars (IV-V) may enhance their crypsis while perching 


VOLUME 38, NUMBER 2 


bs Pe 
_ » tee 


Fic. 5. Fourth instar larva of T. andromeda: dorsal aspects (above); details of the 
head capsule, frontal aspects (below). 





110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 6. Fifth instar larva of T. andromeda. Various aspects of perching and feeding 
behavior (top two photos); general body profile, head capsule, and silk mat perching 
(lower two photos). 


VOLUME 38, NUMBER 2 Aa 





Fic. 7. Pupation and eclosion in T. andromeda. Above: prepupal position and pupa. 
Below: freshly-eclosed adult perched on the empty pupal shell. 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on grass stems. When gently prodded with forceps, larvae of all instars 
quickly regurgitate a dark green fluid from their mouthparts; presum- 
ably, this fluid is a mixture of digestive fluids and partly-digested food 
plant tissues. 


DISCUSSION 


The observations on early stages and food plant association of T. 
andromeda presented here generally agree with what is already well 
documented for other satyrids, including the euptychiines to which 
this genus belongs. The satyrids in general, for example, are associated 
with monocotyledons as larval food plants (Ehrlich & Raven, 1965). 
The observed occurrence of T. andromeda in both old secondary forest 
understory and grassy areas in cacao plantations in eastern Costa Rica 
point to a butterfly that is already documented as being associated with 
a variety of tropical habitats (Ross, 1964, 1976). Although I report only 
one food plant species in this paper, the widespread occurrence of T. 
andromeda throughout Central America (e.g., Monroe et al., 1967) 
suggests that other grasses might also be utilized as larval food planis 
by this butterfly. , 

The description of the first instar larval head capsule for T. ypthima 
by Muller (1886) agrees very well with my description for T. androm- 
eda (Muller’s fig. 28a, b in Plate 13). Other descriptions of instars in 
Muller (1886) generally agree with my findings for T. andromeda. 

While most satyrids are diurnally-active butterflies, some Neotropi- 
cal forms such as Taygetis are crepuscularly-active, and possibly noc- 
turnal. Taygetis is commonly found at rotting fruits in lowland tropical 
wet forest in Costa Rica near dusk (Young, 1972). With the exception 
of a few species, most Taygetis are believed to rely upon olfactory and 
tactile courtship signals, since wing color patterns are very similar be- 
tween the sexes (Forbes, 1952). Certainly, the subdued brown color- 
ation of this large butterfly and the cryptic appearance and behavior 
of the early stages together suggest an insect that is probably quite 
palatable to potential predators, including arthropods and small ver- 
tebrates such as lizards. 

While the early stages of T. andromeda are similar to those of most 
other described satyrids (many papers in this journal for both temper- 
ate and tropical forms), they exhibit some differences which warrant 
further detailed consideration, particularly for comparisons with other 
species of the genus. 


ACKNOWLEDGMENTS 


This research is a by-product of grants from The American Cocoa Research Institute, 
and the Friends of the Milwaukee Public Museum. P. J. DeVries and Lee D. Miller 


VOLUME 38, NUMBER 2 Te 


offered constructive comments on an earlier draft of the manuscript. Tammy McCarthy 
prepared Fig. 4. The comments of two anonymous reviewers were most helpful in 
providing a focus to the discussion section. 


LITERATURE CITED 


D ALMEIDA, R. F. 1922. Melanges Lepidopterologiques. I. Etudes sur les Lepidopteres 
du Bresil. Berlin: R. Friedlander & Sohn, 226 pp. 

EBERT, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of the 
butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soe 23:Suppl. No. 3, 
48 pp. 

EHRLICH, P. R. & P. H. RAVEN. 1965. Butterflies and plants: A study in coevolution. 
Evolution 18:586-608. 

ForRBES, W. T. M. 1952. A draft key to Taygetis (Satyrinae). Lepid. News 6:97-98. 

Lamas, G. 1967. Notas sobre mariposas peruanas (Lepidoptera). III. Sobre una colec- 
cion efectuada en el departamento de Tumbes. Rev. Peru. Entomol. 19:8-12. 

MONROE, R. S., G. N. Ross & R. N. WILLIAMS. 1967. A report on two recent collections 
of butterflies from Honduras. J. Lepid. Soc. 21:185-197. 

MULLER, W. 1886. Sudamerikanische Nymphalidenraupen. Zool. Jahrb. I:417-678. 

Ross, G. N. 1964. An annotated list of butterflies collected in British Honduras in 1961. 
J. Lepid. Soc. 18:11-26. 

1976. An ecological study of the butterflies of Sierra de Tuxtla in Veracruz, 
Mexico (continued). J. Res. Lepid. 15:41-60. 

YOUNG, A. M. 1972. Community ecology of some tropical rain forest butterflies. Amer. 
Midl. Nat. 87:146-157. 

1982. Notes on the natural history of Morpho granadensis polybaptus Butler 

(Lepidoptera: Nymphalidae: Morphinae), and its relation to that of Morpho peleides 

limpida Butler. J. New York Entomol. Soc. 90:35-54. 








Journal of the Lepidopterists’ Society 
38(2), 1984, 114-123 


THE LIFE-HISTORY OF ACTIAS MAENAS 
(SATURNIIDAE)! 


WOLFGANG A. NASSIG 


Arbeitsgruppe Okologie, Zoologisches Institut 
der J. W. Goethe-Universitat, Siesmayerstrasse 70, 
D-6000 Frankfurt am Main, Federal Republic of Germany 


AND 


RICHARD STEVEN PEIGLER? 
303 Shannon Drive, Greenville, South Carolina 29615 


ABSTRACT. Broods of the Southeast Asian Actias maenas Doubleday (=A. leto) 
were reared in Germany and South Carolina utilizing stock from West Malaysia and 
northern Sumatra. Larvae preferred Liquidambar styraciflua and Rhus spp. among a 
variety of hostplants offered. Larval development at 23-28°C required 31 to 40 days; the 
pupal stage lasted 12 to 15 days. The first instar larva is orange with a black head and 
black marking on the tergum. The mature larva (fifth instar) is dark lime green with a 
brown head and green spiny scoli, with yellow bands on the posterior edge of abdominal 
segments 2-7. Females fly prior to mating. Mating commences 1-2 h before sunrise and 
lasts only a few hours. The species appears to be polyvoltine, without pupal diapause. 
Some larvae were killed by a disease caused by the microsporidian Nosema. 


ZUSAMMENFASSUNG. Die siidostasiatische Actias maenas Doubleday (=A. leto) 
wurde in Deutschland und South Carolina (U.S.A.) geziichtet; das Zuchtmaterial stammte 
aus West Malaysia und Nordsumatra. Die Raupen bevorzugten Liquidambar styraciflua 
und Rhus spp. unter den angebotenen Futterpflanzen. Die larvale Entwicklungszeit bei 
23-28°C dauerte 31 bis 40 Tage, das Puppenstadium 12 bis 15 Tage. Das erste Raupen- 
stadium ist orange mit schwarzen Kopf und schwarzer Zeichnung. Die ausgewachsene 
Raupe (5. Stadium) ist dunkel gelbgriin mit braunem Kopf und grossen griinen dornigen 
Sternwarzen, am Hinterrand der Abdominalsegmente 2 bis 7 mit gelben Streifen. Die 
Weibchen fliegen vor der Begattung. Die Paarung findet 1-2 Stunden vor Sonnenaufgang 
statt und dauert nur wenige Stunden. Die Art diirfte polyvoltin sein, ohne Puppendia- 
pause. Etliche Raupen wurden durch Darmkrankheiten getétet, ausgelést durch Nosema- 
Erreger. 


Actias maenas Doubleday (1847) has been known for well over a 
century and is very popular with collectors, yet the larval stages of this 
moth are not well known. The species ranges from sub-Himalayan 
regions of northeastern India through most of the Southeast Asian 
mainland and on the Greater Sunda Islands, a distribution of more 
than 4000 km. The biotope covers diverse biomes including tropical 
rainforest, paratropical rainforest, and notophyllous broad-leaved ev- 
ergreen forest (Wolfe, 1979). 

Adults exhibit striking sexual dimorphism; males (Fig. 1) are bright 
yellow with brown markings, whereas females (Fig. 2) are light green. 


''No. 4 of the series “Contributions to the knowledge of the Saturniidae” by the senior author. 
* Museum Associate in Entomology, Natural History Museum of Los Angeles County. 


VOLUME 38, NUMBER 2 Lid 





Fics. 1-4. Actias maenas. 1, male, live specimen in natural repose; 2, female, pinned 
specimen; 3, first instar larva just prior to molting into second instar; 4, mature larva 
feeding on sweetgum. 


The insect was recently characterized morphologically by Arora and 
Gupta (1979), including drawings and a detailed description of the 
male genitalia and wing venation. Narang and Gupta (1981) reported 
on cytological investigations of this saturniid. Despite these recent stud- 
ies, a considerable amount of taxonomic confusion persists with Actias 
maenas. For examples, (1) the junior synonym leto (Doubleday) con- 
tinues to appear commonly on lists of dealers and labels of specimens; 
(2) the genus has been excessively split, with even the recent Indian 
authors cited above using the generic name Sonthonnaxia Watson, 
whereas other authors (Bouvier, 1936; Allen, 1981; Barlow, 1982) con- 
sidered this large Asiatic species to belong to the African genus Argema 
Wallengren; (3) the so-called insular subspecies or forms appear to us 
to be either distinct species (e.g., isis Sonthonnax from Celebes, and 
ignescens Moore from the Andamans) or simply subjective synonyms 
of A. maenas (e.g., saja van Eecke from Java and Sumatra, and recta 
Bouvier from Sumatra). Satisfactory resolution of these problems must 
await a modern revision of the genus. 

To date virtually no comparative studies have been published on the 


116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pre-imaginal instars within the genus Actias. The early stages of A. 
maenas were described by Roepke (1918), and Packard (1914, pl. 96) 
figured the pupa and cocoons. Gardiner (1982, pl. 8) published a color 
photograph of a mature larva. In the present paper the structure and 
behavior of the pre-imaginal stages are described in greater detail and 
compared with related species. Moreover, we present information on 
courtship behavior, hostplant relationships, diseases, and parasitism. 


Rearing Observations 


In May 1982 the senior author received six cocoons of this species 
from Tapah, Perak, West Malaysia. With adults emerging from these 
cocoons, a mating was achieved and part of the eggs was sent to the 
junior author. Rearing was successful in both Germany and South Car- 
olina. Additionally, another brood was reared by the senior author 
during the winter of 1982-1983 utilizing stock from northern Sumatra, 
Indonesia. Observations and descriptions below are based on the Ma- 
laysian material except where otherwise noted. 

Emergence of the adults was in the evening in Germany, within a 
few hours after sunset. In South Carolina, some adults also emerged 
within 2 hours after sunset, while others of both sexes emerged between 
0200 and 0400 h (Eastern Standard Time). Adults have minimal dif- 
ficulty pushing out of the flimsy cocoons. Wing expansion is complete 
within 45 min after emergence. During expansion the tails do not begin 
to elongate until after the forewings have completely expanded. Both 
sexes are easily excitable, flapping vigorously on the ground or bottom 
of the cage after being disturbed. The long tails on live specimens are 
surprisingly flexible and are not easily broken off. In natural resting 
position these tails are held parallel or nearly so (see Fig. 1), while tails 
of specimens of Argema are frequently crossed in repose. On a vertical 
substrate the moths rest at an angle, i.e., the longitudinal axis of the 
body is positioned parallel against the substrate, but is off 20° or more 
from the vertical, either to the right or the left. 

Females fly after wings have hardened but before emitting phero- 
mone, generally shortly after sunset. This character is an unusual one 
among Saturniidae where females of most species emit pheromone and 
mate prior to their first flight. However, this character is probably 
normal for the Actias group in general; the junior author observed this 
in A. luna (L.) in nature, and Marten (1955) reported the same be- 
havior with Graellsia isabellae (Graélls) in Spain. Mating probably 
occurs in treetops with these insects. Several hours after the virgin 
flight, females emit pheromone. In Germany, the first pairing occurred 
between 0100 and 0300 h (MEZ)? in a cage outdoors. Conditions during 


* mitteleuropdische Zeit—Central European Time. 


VOLUME 38, NUMBER 2 a4 


mating were as follows: cloudless, temperature 15.0°C, relative humid- 
ity 72%. The pair remained united until after 0700 h. In South Carolina 
in early August, an F, pairing occurred between 0430 and 0500 h 
(EST), and lasted until ca. 0800 h. The different mating times observed 
in Germany and South Carolina actually agree in terms of their relation 
to respective times of sunrise. 

Both authors reared larger numbers of Actias sinensis heterogyna 
Mell alongside the broods of A. maenas, and both species demonstrated 
the same respective mating times in Germany and South Carolina, i.e., 
covering the two hours on either side of sunrise. Attempts to hybridize 
A. maenas with the much smaller A. sinensis heterogyna were not 
successful. We were unable to hand-pair either species (including inter- 
and intraspecific matings). In South Carolina adults of both species 
were present in one large cage, and females of both species emitted 
pheromone simultaneously, but males of both species mated only with 
conspecific females. 

The original female in Germany deposited ca. 170 ova during the 
first two nights, and laid only ca. 30 eggs total during the following 
three nights. Most of the last deposited eggs were infertile, as is normal 
for Saturniidae (cf. Miller & Cooper, 1977). Incomplete data on fecun- 
dity recorded for an F, female in South Carolina were very similar to 
the above. In both cases females were killed with a small number of 
eggs remaining in their abdomens in order to preserve the good con- 
dition of these specimens for collections. 

Hostplants in nature given by Arora and Gupta (1979) are Turpinia 
sphaerocarpa Hassk. (=T. pomifera), Staphyleaceae, and Schima wal- 
lichii (DC.) Korth., Theaceae.* Barlow (1982) cited Averrhoa bilimbi 
L., Oxalidaceae, as a host. A specimen from Bogor, Java, in the Rijks- 
museum van Natuurlijke Historie (Leiden, Netherlands) was reared on 
Adinandra dumosa Jack, Theaceae. Roepke (1918) reared his material 
in Java on Canarium, Burseraceae. The moth has also been reared on 
Eucalyptus gunnii Hooker, Myrtaceae (Gardiner, 1982). Since none 
of these plants were available to us, several alternatives were offered 
to the larvae. The food which appeared to be most preferred was 
sweetgum (Liquidambar styraciflua L., Hammelidaceae), although 
larvae in all instars freely accepted staghorn sumac? (Rhus typhina L., 
Anacardiaceae) in Germany, and winged sumac (R. copallina L.), 
smooth sumac (R. glabra L.), and poison ivy (R. radicans L.) in South 
Carolina. Several species of oak (Quercus, Fagaceae) were accepted as 
well, the evergreen species of which are probably the best choice for 
rearing during the northern winter. First instar larvae also fed reluc- 


‘ The spelling, synonymy, and family-classification of plants listed follow the work of Backer and Bakhuizen van den 
Brink (1963-1968). 
5 The North American trees sweetgum and staghorn sumac are grown as ornamentals in Germany 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Development of F, brood of Actias maenas on sweetgum in Germany (23- 
28°C) and South Carolina (25-28°C). Mating occurred 31 May 1982. 


Eclosion Molt 1 Molt 2 Molt 3 Molt 4 Spinning 
Germany 10-14 June 15-18 20-24 26-30 2-6 July 17-23 
South Carolina 12-13 June 17-18 22- — 28-30 2-5 July 13-17 


tantly on rose (Rosa, Rosaceae), black walnut (Juglans nigra L., Jug- 
landaceae), and Staphylea colchica Stev., Staphyleaceae, although both 
authors reared larvae mainly on sweetgum. Some of these other plants 
might be utilized successfully by rearers where sweetgum and sumac 
are not available. Actias maenas has been reared in the Amsterdam 
Zoological Garden (van Eecke, 1913; W. Hogenes, pers. comm.), but 
no records of which hostplants were used could be located. 

The hostplants accepted in nature and captivity by Actias maenas 
coincide with those known for related species. Argema mimosae (Bois- 
duval) from Africa feeds on walnut and certain Burseraceae and An- 
acardiaceae (Pinhey, 1972), and Argema mittrei (Guérin-Méneville) 
from Madagascar accepts several species of Rhus (Villiard, 1969) and 
certain Myrtaceae (Pinhey, 1972). Actias sinensis heterogyna from 
Taiwan, partially sympatric with A. maenas, feeds on Liquidambar 
formosana Hance in nature (Mell, 1950) and L. styraciflua in captivity 
(Nassig, 1980). Walnut, oak, and sweetgum are well-known hosts of 
the Nearctic Actias luna, and are accepted by the Asiatic A. selene 
(Hiibner) as well. Sweetgum is also used in nature in Mexico by Actias 
truncatipennis Sonthonnax (Beutelspacher, 1978). 

Newly hatched larvae are very active and move quickly about in 
the rearing container. Larvae show minimal tendency to congregate, 
even in the first instar. The resting position is with the anterior end of 
the body held free of the leaf or stem, the typical “sphinx” position for 
Sphingidae and most Saturniidae. Larval development was rapid under 
temperatures of 23-28°C, the time from eclosion of eggs to spinning 
of cocoons being ca. 31 to 40 days (Table 1). Males complete devel- 
opment a few days more rapidly than sibling females, both in the larval 
and pupal stages, because they are smaller. Equally good results were 
achieved by rearing in open air (Ndssig) and under plastic bags (Peig- 
ler) to ensure high relative humidity; both of us reared all larvae in- 
doors on cut food with stems inserted into water. 

Structurally, larvae of A. maenas are similar to other species of 
Actias and Argema. The first instar is orange as in A. selene from 
Bhutan, A. sinensis heterogyna, A. artemis? (Bremer) from South Ko- 
rea, and Argema mimosae (only A. luna is green in the first instar). 


VOLUME 38, NUMBER 2 119 


In the second instar the body of A. maenas becomes yellowish green 
as in the case of A. artemis?; A. selene and A. sinensis heterogyna 
remain orange. The third instar is green for all species. The subspirac- 
ular yellow stripe of the mature larva of A. selene and A. artemis? is 
absent in Argema and all other species of Actias mentioned above. 
Mature larvae of A. maenas look much more like those of Argema 
than any Actias which we have seen. The dark green integument with 
contrasting short whitish setae are seen in Argema mittrei (Villiard, 
1969), while the elongated fleshy extensions of the body which support 
the dorsal scoli are also present in Argema mimosae (Pinhey, 1972:pl. 
1) and to a lesser degree in A. selene. All species of Actias and Argema 
for which the larvae are known have a single median dorsal scolus on 
abdominal segment 8, a character which easily separates the group 
from several similarly appearing larvae of Saturnia Schrank and allied 
genera in which a pair of dorsal scoli is present on abdominal seg- 
ment 8. 

In the later instars of Actias maenas, the scoli bear stout spines, 
especially the dorsal scoli. These possibly have mechanical defensive 
benefits, because they lack fluid secretions seen in several other satur- 
niid caterpillars, e.g., Saturnia pyri [Denis & Schiffermiiller] (Haffer, 
1921). In the last three instars the ventral side is a darker green than 
the dorsal side, this providing a very effective camouflage known as 
countershading (de Ruiter, 1955). 

Dupont and Scheepmaker [1986] stated that larvae of Actias reared 
in Java were very susceptible to disease, a problem also noted by Roepke 
(1918). All larvae of an F, brood in South Carolina succumbed to a 
disease before reaching the last instar. Of the four main types of patho- 
gens infecting Saturniidae, viz. viruses, bacteria, fungi, and microspo- 
ridians, outlined by Jolly et al. (1979:61-66), the symptoms agreed with 
the latter. The pathogen belongs to the genus Nosema (Nosematidae). 
The symptoms include reduced feeding, lethargy, black spots on the 
integument, darker body color, and stunted growth. A large portion of 
the F, brood of larvae in Germany was also killed by an unidentified 
disease. Higher humidity is beneficial to Lepidoptera which are native 
to tropical rainforests, but unfortunately such conditions also promote 
disease. 

Cocoons of Actias maenas are attacked in nature by one or more 
species of Xanthopimpla (Hymenoptera: Ichneumonidae) belonging to 
the regina species-group (Townes & Chiu, 1970). We were unable to 
locate other published records of parasitism for this moth. 

Mature larvae spin cocoons among leaves of the hostplant or among 
dead leaves on the bottom of the cage. There appeared to be no pref- 
erence in selection of a pupation site. In nature, presumably some 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Behavioral characteristics of Actias maenas considered to be of generic 
level. 


First instar larvae move about rapidly 

Larvae not gregarious, even in first instar 

Pupa with spines on cremaster anchored into cocoon 

Pupa frequently active 

Females fly prior to emitting pheromone 

Adults excitable at all times; copula is broken at slightest disturbance 
Adults rest at angle on vertical substrate 

Predilection for Liquidambar and Anacardiaceae as hostplants 


cocoons are formed at ground level while others are spun high above 
the ground among living leaves. Freshly spun cocoons are white; the 
silk usually then turns light or dark brown. Most cocoons have small 
perforations as in the cocoons of Actias selene, A. sinensis heterogyna, 
and Argema mimosae. A few cocoons of A. maenas have less than five 
discernable perforations and occasionally none at all. Packard (1914) 
figured an imperforate cocoon alongside one with numerous perfora- 
tions. The imperforate cocoon was also stated to be a ‘valveless’ cocoon, 
but we have not seen cocoons of A. maenas which lack the pre-formed 
exit at the anterior end. The hooked spines of the pupal cremaster are 
anchored into the posterior end of the cocoon. This undoubtedly fa- 
cilitates the emerging moth when pulling itself out of the pupal shell. 
The active pupa is frequently heard rolling in the papery cocoon at 
any time during the pupal stage, especially when disturbed by an 
external stimulus (refer to Table 2). 

Diapause of some species of Actias has been investigated. In general, 
photoperiod is the primary mechanism which maintains and termi- 
nates diapause in temperate species such as A. luna (Wright, 1970) 
and Japanese species (Miyata, 1974, 1976, 1977). Actias maenas is a 
tropical species however, the range of which crosses the Equator. Ac- 
tias sinensis heterogyna, with a slightly more northern range, dia- 
pauses only rarely: only one female among ca. 100 pupae in Germany 
diapaused for six months at 6°C; short-day photoperiod, senescing host- 
plants, and lower temperatures during larval and pupal stages all failed 
to induce diapause in A. s. heterogyna (Nassig, 1980 and unpubl.; Mell, 
1950). These data plus the rapid life cycles indicate that both A. mae- 
nas and A. s. heterogyna are polyvoltine in nature, notwithstanding 
the presence of a light-detecting “window” on the head of the pupa in 
each species (see Fig. 5). However, Bouvier (1936:254) indicated that 
A. maenas is bivoltine in northeastern India, a region where mild 


VOLUME 38, NUMBER 2 LAA 





ANNAN; Si 


ti iy ie 






Fic. 5. Actias maenas, ventral view of male pupa. Note transparent “window” be- 
tween eyes and base of antennae, and strands of silk from cocoon attached to cremaster. 


winters occur (Wolfe, 1979). No seasonal forms are known for adults 
of either species. 


Descriptions of Immature Stages 


The following descriptions are based on living material, freeze-dried 
larvae, and color transparencies. The minor differences noted between 
material from West Malaysia and Sumatra are not believed to be tax- 
onomically significant. The photographs and drawing in this paper 
were all made by the senior author. 


Ovum. Length 2.5 mm, width 2.1 mm, height 1.6 mm. (Sizes of eggs from Malaysian 
and Sumatran material are the same, but Roepke (1918) cited smaller measurements for 
Javanese eggs.) Coloration whitish brown. Irregular areas of chorion opaque or translu- 
cent. Chorionic sculpturing evenly reticulate, average diameter of meshes 0.02 mm. 
Partially coated with brown secretion for affixing egg to substrate. 

Larva. First instar (Fig. 3). Head glossy black, 1.1 mm in diameter. Integument 
orange; dorsal and lateral area of abdominal segments 1 to 4 black. Thoracic legs, prolegs, 
and posterior edge of anal plate all black. Scheme of implantation of scoli in six longi- 
tudinal rows, with two rows each of dorsal, subdorsal, and lateral, excepting single median 
dorsal scolus on abdominal segment 8. (Roepke’s (1918) statement that there are only 
four rows of scoli we believe to be erroneous.) Scoli not prominent, concolorous with 
integument, each with 4-6 short white primary setae, longer ones on thoracic and sub- 
spiracular scoli. Larval length reaching more than 1 cm. 

Second instar. Head glossy dark brown, lighter frons and clypeus, ca. 2.5 mm in 
diameter. Integument light yellowish green, with irregular tiny white granulations, each 
bearing seta. Thoracic legs and dorsal portion of prolegs black to dark brown. Prothoracic 
plate in some individuals with an irregular black patch, more prominent in Sumatran 
material. Anal plate dark brown with yellow border. Subspiracular line yellow, connect- 
ing subspiracular scoli, less distinct in Sumatran material. Bases of scoli yellow, distally 
becoming orange or red in dorsal and subspiracular rows, yellow in lateral rows. Each 
scolus with single black seta arising from center, and 4-7 radial black spines. Larval 
length reaching 2 cm. 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Third instar. Head glossy brown, lighter frons and clypeus, 3.5 mm in diameter. 
Integument dark lime green, with numerous contrasting white granulations, each with 
seta. Thoracic legs dark brown. Prolegs dark gray. Prothoracic plate green, concolorous 
with integument. Anal plate light brown with ca. 20 small black spots, anterior edge 
yellow; anal prolegs dark brown with anterior edge yellow. Spiracles green, concolorous 
with integument. Subspiracular stripe yellow. Scoli yellow, becoming green distally, in 
Sumatran material sometimes all but largest scoli remaining yellow or even orangish with 
basal red ring, as in other species of Actias. Larval length ca. 3.2 cm. 

Fourth instar. Same as third instar except head 4.0—4.5 mm in diameter, subspiracular 
stripe disappearing, scoli all becoming larger, and fleshy extensions of body supporting 
scoli becoming more prominent. Larval length ca. 6 cm. 

Fifth instar (Fig. 4). Same as third and fourth instars except head 5-7 mm in diameter, 
spiracles dark grayish yellow, scoli each with 4-10 central and radial black spines 0.5— 
1.5 mm long, best developed on dorsal pairs of meso- and metathoracic scoli plus median 
caudal scolus (this scolus was lost on specimen figured due to an accident), and posterior 
edge of first seven abdominal segments with light yellow stripe. Larval length 9.5-10.5 
cm. 
Pupa (Fig. 5). Color dark brown. Antennal covers small, surrounded by leg covers. 
Head with transparent (whitish yellow) “window” between compound eye covers. Ab- 
dominal segments telescoped to small degree, pair of small protuberances on ventral side 
of segments 5 and 6 homologous to prolegs (cf. Mosher, 1916). Cremaster with several 
hooked spines. Length of male ca. 4 cm, female ca. 5 cm, width 1.5-2.0 cm (reared 
material being smaller than this). 

Cocoon. Ovoid, irregular in shape. Length 5-6 cm, width 2.0-3.5 cm. Texture papery, 
spun between leaves. Coloration light brown with glossy sheen. Usually sparsely perfo- 
rated. Pre-formed exit opening present at anterior end. Occasionally incomplete net-like 
inner cocoon is observed. 


ACKNOWLEDGMENTS 


We are grateful to Dr. Rienk de Jong (Rijksmuseum van Natuurlijke Historie) and 
Willem Hogenes (Instituut voor Taxonomische Zoélogie, Amsterdam) for providing key 
literature references and searching for hostplant records among the long series of pinned 
Actias maenas in their respective museums. We wish to thank Dr. Dieterich, Meteoro- 
logisches Institut der J. W. Goethe-Universitat, Frankfurt, for the weather data. Ian 
Wallace of Halesowen, England, supplied livestock of A. maenas. 


LITERATURE CITED 


ALLEN, M. G. 1981. The saturniid moths of Borneo with special reference to Brunei. 
Brunei Mus. J. 5:100-126. 

ARORA, G. S. & I. J. Gupta. 1979. Taxonomic studies on some of the Indian non- 
mulberry silkmoths (Lepidoptera: Saturniidae: Saturniinae). Mem. Zool. Surv. India 
16. ii + 63 pp., 11 pls. 

BACKER, C. A. & R. C. BAKHUIZEN VAN DEN BRINK. 1963-1968. Flora of Java (Sper- 
matophytes only), vols. 1-3. Wolters-Noordhoff, Groningen. 

BARLOW, H.S. 1982 [1983]. An introduction to the moths of South East Asia. Malayan 
Nature Soc., Kuala Lumpur. ix + 305 pp., 50 col. pls. 
BEUTELSPACHER, C. R. 1978. Familias Sphingidae y Saturniidae (Lepidoptera) de Las 
Minas, Veracruz, México. An. Inst. Biol. Univ. Auton. (Ser. Zool.) 491:219-229. 
BOUVIER, E.-L. 1936. Etude des Saturnioides normaux, famille des Saturniidés. Mém. 
Mus. Natl. Hist. Nat., Paris (n. sér. 3):1-354, figs. 1-82, pls. 1-12. 

DOUBLEDAY, E. 1847. Description of a new species of the genus Actias of Hubner, 
from northern India. Ann. Mag. Nat. Hist. 19:95, pl. 7. 

DUPONT, F. & G. J. SCHEEPMAKER. [1936]. Uit Java’s vlinderleven. Visser & Co., Ba- 
tavia. 216 pp. 


VOLUME 38, NUMBER 2 123 


VAN EECKE, R. 1918. On the varieties of Actias maenas, Doubld. Notes Leyden Mus. 
35:132-139, pls. 3-6. 

GARDINER, B. O. C. 1982. A silkmoth rearer’s handbook. Amat. Entomol. 12, xiii + 
255 pp., 26 +[8] h.-t. pls., 32 col. pls. 

HaFFER, O. 1921. Bau und Funktion der Sternwarzen von Saturnia pyri Schiff. und 
die Haarentwicklung der Saturniidenraupen. Arch. Naturgesch. (A)87:110-166. 
JOLLY, M. S., S. K. SEN, T. N. SONWALKAR & G. K. PRASAD. 1979. Non-mulberry silks. 

Food & Agric. Org. United Nations, Serv. Bull. 29, Rome. xvii + 178 pp. 

MARTEN, W. 1955. Ueber die Lebensgeschichte von Graéllsia isabellae (Grls.) nebst 
Beschreibung einer neuen Varietat dieser Art. Entomol. Z. (Stuttgart) 65:145-157. 

MELL, R. 1950. Aus der Biologie der chineschen Actias Leach. Entomol. Z. (Stuttgart) 
60:41-—45, 53-56. 

MILLER, T. A. & W. J. COOPER. 1977. Oviposition behavior of colonized Callosamia 
promethea (Saturniidae). J. Lepid. Soc. 31:282-283. 

MiyATA, T. 1974. Studies on diapause in Actias moths (Lepidoptera, Saturniidae). I. 
Photoperiod induction and termination. Kontya 42:51-63. 

1976. Studies on diapause in Actias moths II. The sensitive stage of photoperiod, 

threshold temperature and thermal constants for development with seasonal life 

cycle in different parts of Japan (Lepidoptera: Saturniidae). Trans. Lepid. Soc. Jap. 

26:103-109 [in Japanese with English summary]. 

1977. Studies on diapause in Actias moths (Lepidoptera, Saturniidae). III. 
Effects of photoperiod and temperature on the occurrence of seasonal forms. Kontyt 
45:320-329 [in Japanese with English summary]. 

MOosHER, E. 1916. The classification of the pupae of the Saturniidae. Ann. Entomol. 
Soc. Amer. 9:136-156, pls. 5-6. 

NARANG, R. C. & M. L. Gupta. 1981. Chromosome studies including a report of B- 
chromosome in a wild silkmoth, Sonthonnaxia maenas (Doubleday) (Saturniidae: 
Saturniinae). J. Res. Lepid. 18:208-211. 

NAssic, W. 1980. Zur Zucht von Actias sinensis Walker (Attacidae). Nachr. Entomol. 
Ver. Apollo, Frankfurt 4:42—48. 

PACKARD, A. S. 1914. Monograph of the bombycine moths of North America, part 3 
(T. D. A. Cockerell, ed.). Mem. Natl. Acad. Sci. 12:ix + 1-276 + 5038-516, 113 pls. 

PINHEY, E. 1972. Emperor Moths of South and South Central Africa. C. Struik, Cape 
Town. xi + 150 pp., 48 pls. 

ROEPKE, W. 1918. Over het opkweeken van eenige merkwaardige vlindersoorten uit 
eieren. I. Actias maenas. De Tropische Natuur 7:116—122. 

DE RUITER, L. 1955. Countershading in caterpillars. Arch. Néerl. Zool. 11:285-341. 

TowngEs, H. & S.-C. Curu. 1970. The Indo-Australian species of Xanthopimpla (Ich- 
neumonidae). Mem. Amer. Entomol. Inst. 14:iii + 372 pp. 

VILLIARD, P. 1969. Moths and How to Rear Them. Funk & Wagnalls, New York. xiii 
+ 242 pp. 

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Geol. Surv. Prof. Paper 1106. iii + 37 pp., 3 col. pls. 

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in the wild silkworm, Actias luna. J. Lepid. Soc. 24:209-212. 








Journal of the Lepidopterists’ Society 
38(2), 1984, 124-133 


MYRMECOPHILY IN THE EDWARD’S HAIRSTREAK 
BUTTERFLY SATYRIUM EDWARDSII (LYCAENIDAE) 


R. P. WEBSTER! AND M. C. NIELSEN 


Department of Entomology, Michigan State University, 
East Lansing, Michigan 48824 


ABSTRACT. Observations on the life history and myrmecophilous relationship of 
the lycaenid butterfly, Satyriwm edwardsii (Grote and Robinson) and the ant, Formica 
integra Nylander, are described. S. edwardsii departs from other North American Ly- 
caenidae in that the 3rd and 4th instar larvae aggregate during the day (in groups of up 
to 114 individuals per host) at the base of the host plant (Quercus velutina and Q. 
coccinea saplings) within conical structures of detritus (byres) constructed by the ants. 
The larvae leave the byres at dusk, feed nocturnally, and are usually surrounded by a 
group of attending ants. A membracid, Similia camelus (Fabricius), was abundant on 
the same host plants and was ant attended. We suggest that membracids associated with 
S. edwardsii larvae may be involved in the symbiotic relationship between S. edwardsii 
and F. integra. 


Many species of Lycaenidae in North America are myrmecophilous 
(Downey, 1961, 1962; Harvey, 1980). The larvae are commonly sur- 
rounded by a group of ants that groom and palpate them with their 
antennae. On the 7th abdominal segment of late instar larvae is a dorsal 
gland, called Newcomer’s organ, that secretes honeydew on which the 
ants feed (Newcomer, 1911; Malicky, 1970; Maschwitz et al., 1975). In 
addition, there are epidermal glands which secrete substances that at- 
tract and appease the ants (Malicky, 1970). In Glaucopsyche lygdamus 
(Doubleday) these attractive substances secure ant defense against 
parasitoid attack. This protection probably acts as a potent selective 
force in maintaining the symbiosis between lycaenid larvae and ants 
(Pierce & Mead, 1981). 

The symbiotic relationship between ants and lycaenid larvae has 
received comparatively little attention for North American species, and 
in only a few cases have the associated ants been identified. Clark 
(1932) mentioned an association of Satyrium edwardsii (Grote & Rob- 
inson) with ants. Later Comstock (1940) reported that the eggs over- 
winter and that the larvae could be found in “‘ant nests’’ at the roots of 
scrub and scarlet oaks. Other than a brief larval description by Scudder 
(1889), little additional information is available on the biology of S. 
edwardsii. We here report on some observations of the life history and 
myrmecophilous relationship of S. edwardsii and the ant, Formica 
integra Nylander, in Michigan. It is our hope that this preliminary 
and somewhat anecdotal account will stimulate further studies on the 
myrmecophilous relationships between lycaenids and ants. 


‘Current address: Department of Entomology, University of Massachusetts, Amherst, MA 01003. 


VOLUME 38, NUMBER 2 LZ 


STUDY AREAS AND METHODS 


S. edwardsii has been reported from several localities in the lower 
peninsula of Michigan (Moore, 1960). We found this hairstreak to be 
abundant in the Flat River State Game Area, Montcalm Co., TON, 
R7W, Sections 29-30 (Locality 1), and in Newaygo Co., T12N, R12W, 
Sections 1-2 (Locality 2). In these localities the primary host plants of 
S. edwardsii were black oak, Quercus velutina Lam. and scarlet oak, 
QO. coccinea Muench. A few larvae were also found on a Q. alba L. 
sapling in locality 2. Populations of S. edwardsii appeared to be closely 
associated with colonies of F. integra. Adjacent localities without F. 
integra did not support populations of S. edwardsii even though suit- 
able host plants were present. Most of our observations were made at 
locality 1 in Montcalm Co. at various intervals (usually once a week) 
from April through July during 1980 and 1981. 

Locality 1. The study area was a second growth woodlot of Q. 
velutina, QO. coccinea and OQ. alba with a mixture of Populus tremu- 
loides Michx., Pinus strobus L., and miscellaneus upland hardwood. 
S. edwardsii adults were abundant during July and occurred most 
frequently within a narrow, irregularly shaped opening of approxi- 
mately 0.6 hectare. Within the opening were small QO. velutina and 
QO. coccinea with base diameters between 1.3 and 10.0 cm (most be- 
tween 2.5 and 5.0 cm). Additional plants in the openings were Lupinus 
perennis L., Ceanothus americanus L., Rudbeckia hirta L. and as- 
sorted grasses and forbs. C. americanus appeared to be the major 
nectar source for the adults. The soil consists of Graying Sand (Schnei- 
der, 1960), a deep sand with low waterholding capacity. Five large 
(0.8-1.2 m dia.) nests of F. integra were in the opening. 

Locality 2. This colony was located at the edge of a relict prairie. 
The same species composition of trees was found as at locality 1. Q. 
velutina saplings were the dominant species within the openings scat- 
tered within the second growth forest. Few other plants occurred in 
the openings other than grasses, forbs, and scattered clumps of C. 
americanus. The F. integra nests were smaller (no well defined mounds) 
and were situated within clearings. The soil was a Grayling Sand. 


OBSERVATIONS 


Larval, ant, and membracid associations. The eggs of S. edwardsii 
hatched during late April and early May when the oak buds were 
enlarged and ready to open. The reddish brown Ist instar larvae bored 
into the buds of Q. velutina and Q. coccinea saplings and suckers and 
fed diurnally. The 2nd instar larvae continued to feed on the buds and 
developing leaves. Buds with Ist or 2nd instar larvae usually had 2-4 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


larvae, but sometimes 6 or 7 were present. The larvae were frequently 
near nymphal aggregations of the membracid, Similia camelus (Fa- 
bricius) and scale insects (Coccoidea). These Homoptera release hon- 
eydew and were ant attended. The ants, in turn, defended any oak 
sapling having these Homoptera. When a sapling or branch was dis- 
turbed most ants assumed a defensive posture (reared up on pro- and 
mesothoracic legs with mandibles open, and sometimes pointed the 
abdomen anteriorly between the metathoracic legs), frequently mov- 
ing toward the source of the disturbance. The ants readily attempted 
to bite any object brought near them. Although the lst and 2nd instar 
larvae presumably can not produce honeydew (they do not have New- 
comer’s organs), ants were usually within 2.5 cm of each larva and 
appeared to tend them. We found Newcomer’s organs only on 4th 
instar larvae. 

Around the roots at the base of the oak saplings were chambers that 
extended to 7.5 cm below the soil line along the taproot and laterally 
around the roots up to 10 cm away from the taproot. There was a 1- 
3 cm space around the roots and a small amount of debris covering 
the opening at the soil surface. Trenches and tunnels (beneath a loose 
layer of dried grass and leaves), led away from the chambers to ant 
nests that were 1-13 m away. No Ist and 2nd instar larvae of S. 
edwardsii or membracid nymphs were found in the chambers with 
the ants during late April or early May. 

By late May the ants had constructed conical structures (byres) of 
fine pieces of detritus (much like the material covering the ant nests) 
at the bases of the oak saplings (Figs. 1 & 2). The byres were 10 to 25 
cm in dia. and extended 5 to 20 cm upward on the stems of the oaks. 
The largest byres were around the largest saplings (10.0 and 20.0 cm 
dia.). There was a 1.0-1.5 cm space between the inner side of the byre 
and the bark of the sapling confluent with the chambers around the 
roots (Fig. 2). The upper edge of the byre was contiguous with the 
bark of the sapling except for a series of 1-4 mm gaps which allowed 
access of the ants into and out of the byres. 

The behavior of the larvae of S. edwardsii changed when they 
reached the 8rd instar (during late May). Although there was consid- 
erable evidence of feeding, larvae were not found on the foliage during 
the day. Instead, the mottled, brownish 3rd and 4th instar larvae re- 
mained during the day on the bark in the chambers within the byres 
at the base of the oak saplings. Only 4 larvae were found outside the 
byres during the day, and these were on branches of the 10 cm dia. 
sapling among aggregations of S$. camelus nymphs and adults. The 
larvae in the byres were positioned vertically on the stem, at or above 


VOLUME 38, NUMBER 2 


Fic. 1. (Top) A 2.5 cm dia. Q. velutina with a byre (arrow) at base. (Bottom) Six 
Ath instar larvae of S. edwardsii with attending ants and a S. camelus nymph (arrow) 
within a byre at the base of a 1.9 cm dia. Q. velutina. The stem of the oak was pulled 
away from the observer to reveal the larvae in the byre. The membracid nymph is 
positioned where the upper edge of the byre had been. 





128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. A 3.5 cm dia. Q. velutina with a byre at base. A portion of the byre was 
removed to reveal the larvae and space (arrow) between inner side of byre and bark of 
sapling. 


the soil line (Fig. 1). Between 1 and 114 larvae were found in the byres 
and the largest larval aggregations were in those byres on saplings with 
base diameters of 3.8 to 10.0 cm (Table 1). Larvae and byres were not 
found on Q. velutina and Q. coccinea with base diameters greater 
than 20 cm or on other species of shrubs or trees. Between 20 and 100 
(an exact count was difficult) ants were in the chambers with the larvae 
in each of the byres. Only 3 S. camelus were found in the byres, 
although many nymphs and adults (and attending ants) were on the 
branches of the saplings. One 1.3 cm dia. sapling had 138 S. camelus 
nymphs and adults and 75-100 ants on it. The ants appeared to obtain 
honeydew from the membracids. 


VOLUME 38, NUMBER 2 129 


TABLE 1. Number of S. edwardsii larvae in byres at bases of Q. velutina and Q. 
coccinea saplings at locality 1. Observations were made on 8 June 1980 and 6 June 1981. 


Sapling size x no. larvae! x no. larvae! 
(dia. at base in cm) 1980 1981 
een 1.0 (2) 4.5 (2) 
Lg 8.5 (2) — 
2.5 BP 2SS) OC) 
3.1 = 9.0 (1) 
3.8 2.7 (3) — 
5.0 19.7 (8) 20.5 (2) 
6.3 13.0 (1) 22.0 (1) 
10.0 60.0 (1) 2S OV.GR)2 
20.0 3.0 (1) _- 
>20.0 0.0 (8) — 


1 Numbers in ( ) are the number of saplings examined in size class. 
2 Ninety-one larvae (out of 114) were removed from this sapling on 31 May. 


The 8rd and 4th instar larvae of S. edwardsii fed on the foliage of 
the host only at night. They left the byres within one hour after sunset, 
crawled up the stem and fed on the foliage until sunrise, and then 
returned to the byre. On 14 June (1980), 4th instar larvae on a sapling 
(10 cm dia.) and a sucker clump (17 larvae in byre) were observed. A 
dim flashlight was used to observe the insects at night. Sunset on 14 
June was 2117 h EDT and sunrise on 15 June was 0559 h. At 2140 h 
(0.5 h after sunset) one larva left the byre on the 10 cm dia. sapling. 
By 2155 h several more larvae had left the byre and were crawling up 
the trunk. One larva crawled at a rate of 1.8 cm/min. At 2225 h all 
17 larvae on the sucker clump were crawling up the stems. Each larva 
was accompanied by 1-8 ants as it crawled up the stem. At least one 
ant was always within 2.5 cm of each larva, and occasionally an ant 
was observed on the dorsal surface of a larva. By 2300 h the larvae 
had begun to feed on the oak foliage and continued to feed throughout 
the night. Between 0605 and 0630 h (5-30 min after sunrise) the larvae 
began leaving the foliage and crawled down the branches and stems 
of the host. At 0615 h only 8 of the 17 larvae on the sucker clump 
were feeding, the rest had either entered the byres or were crawling 
down the stems. On the 10 cm dia. oak 2 larvae were crawling down 
the main trunk 2 and 3 m above the forest floor, while another larva 
was just entering the byre at the base of the oak. During this time the 
sky was overcast and the temperature was 21°C. No larvae were ob- 
served outside the byres after 0700 h. Throughout our observations 
ants were always on or within 2.5 cm of each larva and appeared to 
obtain honeydew from them. 

On 31 May (1981), 91 larvae (out of 114) were removed from the 
10 cm dia. Q. velutina for determination of parasitism. The larvae 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


were reared to maturity on fresh Q. velutina leaves. Percent parasitism 
was 26% and was due to Tachinidae and Braconidae. It is not known 
which larval stadia are attacked or the mode of parasitoid attack. 

Pupation. Pupation occurred during late June and early July at the 
base of the host plant, either on the underside of leaves that were 
underneath or adjacent to the byre, or on the stem of the host within 
the byre. On 29 June, of the 17 larvae observed on 14 June, 6 had 
pupated on the underside of leaves adjacent to or under the byre, 3 
were on the main stems in the byre (these were parasitized by braconid 
wasps), and 3 groups of braconid cocoons were on the underside of the 
leaves. This accounted for 12 of the 17 observed earlier. 

The pupae of S. edwardsii, like many other lycaenids (Downey, 
1966), produced a faint rattling or creaking sound. The sound was 
produced only after the pupae were disturbed. Because few ants were 
present in the byres (1-4) or on the saplings (less than 20) it remains 
unclear what role (if any) the sounds might play in the myrmecophi- 
lous relationship of S. edwardsii and F. integra. No S. camelus nymphs 
or adults were present, even though adults were abundant on 14 June. 
This might partially account for the paucity of ants on the saplings. 

Adult behavior. One freshly emerged S. edwardsii adult was ob- 
served on 29 June (1981) and by 9 July adults of both sexes were 
numerous. Visual count suggested that between 250 and 350 individ- 
uals were within the clearing. Adults nectared on C. americanus and 
R. hirta. C. americanus was the dominent flowering plant at this time 
and was the major nectar source for the adults. Harkenclenus titus 
(Fabricius), S. liparops (Leconte), and S. calanus (Hiibner) were also 
nectaring at the C. americanus flowers. Males of S. edwardsii were 
usually on leaves at the tops of the larger saplings and shrubs within 
and bordering the clearing. They frequently engaged in aerial ““combat”’ 
with other hairstreaks that flew near them. The ants did not display 
any aggressive behavior towards the adult butterflies. An ant that en- 
countered a female walking on a branch, stopped, palpated her with 
its antennae, and then walked away. 

Oviposition of 2 S. edwardsii females was observed between 1330 
and 1347 h on 9 July (1981). One female walked along a horizontal 
branch of a Q. velutina sapling, probed her abdomen into a knobby 
wound in a small fork, deposited an egg, and then flew away. The 
other female oviposited an egg in a vertical wound about 20 cm above 
the forest floor on a 3.1 cm dia. sapling. The egg was placed under the 
rough bark that formed the edge of the wound. During September, 
several Q. velutina saplings were examined for ova. All ova were either 
in old wounds or hidden under loose bark or dead wood, and occa- 


VOLUME 88, NUMBER 2 131 


sionally, empty egg shells from the previous year were adjacent to the 
newly laid eggs. In one wound 4 ova were stacked on top of each other. 
Most ova were between 0.6 and 1.5 m above the forest floor. 


DISCUSSION 


S. edwardsii is myrmecophilous as are many North American Ly- 
caenidae. However, S. edwardsii departs from its congeners and other 
Lycaenidae in North America in that the 8rd and 4th instar larvae 
aggregate during the day at the base of the host within conical struc- 
tures of detritus constructed by the ant, F. integra. The 8rd and 4th 
instar larvae feed nocturnally on the foliage and are frequently sur- 
rounded by a retinue of ants. Other Satyrium species are tended by 
ants, but they do not form aggregations and usually remain on the host 
leaves or fruit, leaving them only to pupate. 

Fourth instar larvae of S. edwardsii produce honeydew on which 
ants feed. The larval aggregations might, therefore, provide a valuable, 
highly attractive and easily defended food resource for the ants as in 
many Australian and South African Lycaenidae (Clark & Dickson, 
1971; Common & Waterhouse, 1972; Pierce & Mead, 1981). The byres 
should facilitate protection of larvae by ants and provide the ants with 
more ameliorating environmental conditions (higher relative humidi- 
ty). The larvae in turn would gain protection from predator and para- 
sitoid attack. In the riodinid, Anatole rossi Clench, “pens” were con- 
structed by ants at the base of the host plant in response to the presence 
of a larva on the host but only after the honey glands became functional 
(Ross, 1966). However, it is unclear from the present evidence if the 
aggregations of S. edwardsii larvae provide the stimuli that induce 
byre construction by F. integra. The 8rd instar larvae of S. edwardsii 
which do not have functional Newcomer’s organs also rest in the byres. 

The membracid, S. camelus, and other homopterans associated with 
S. edwardsii larvae may play a role in the symbiotic relationship be- 
tween S. edwardsii and F. integra. Membracids release honeydew and 
thus, become an important energy resource for the ants. The ants in 
turn become a resource for the membracids because they reduce pre- 
dation on nymphs (Wood, 1977, 1982b). The effectiveness of ants in 
promoting membracid survival depends on factors such as the number 
of nymphs present, the proximity and size of the ant colony, and the 
longevity of the host plant (McEvoy, 1979; Wood, 1982a, b). Long- 
lived perennials like Q. velutina and Q. coccinea provide relatively 
predictable oviposition sites for female membracids. The nymphal ag- 
gregations that result provide a predictable energy resource for the 
ants. Ant colonies established close to such membracid host plants will 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


promote nymphal survival, as well as increased fitness to the ant colony 
(Wood, 1982a). The presence of nymphal aggregations of S. camelus 
near S. edwardsii larvae may concomitantly benefit the larvae by in- 
creasing the number of ants in their vicinity. This may be particularly 
important for the protection of early instar larvae which are incapable 
of producing an energy resource (honeydew) for the ants. Compounds 
produced by the epidermal glands probably facilitate protection of the 
early instar larvae by keeping those ants already present in their vicin- 
ity. 

The pupae of S. edwardsii produce a faint creaking or rattling sound 
after being disturbed. Downey (1966) reported sound production in 
pupae of six additional Satyrium species, as well as in many other 
lycaenids and three riodinids. The stridulatory organs are located be- 
tween the fifth and sixth abdominal tergites and produce the noise 
when the abdominal segments are rapidly moved. It is not clear what 
the function of the sounds is for S. edwardsii or other Lycaenidae 
(Downey, 1966). In A. rossi the stridulatory organs in conjunction with 
pupal glands on the metathoracic segment appear to serve as ant- 
attractant organs (Ross, 1964, 1966). 

The myrmecophilous relationship between S. edwardsii and F. in- 
tegra is undoubtedly one of the more advanced symbioses known among 
any of the North American Lycaenidae and ants. Additional studies 
are required to ascertain the degree of protection of the larvae offered 
by the ants from predation and parasitism and to detail the precise 
relationships between S. camelus, S. edwardsii, and F. integra. 


ACKNOWLEDGMENTS 


We wish to thank J. C. Nickerson and W. F. Buren, University of Florida/Florida 
Department of Agriculture, Division of Plant Industry, J. Y. Miller, Allyn Museum of 
Entomology, and Dr. A. Francoeur, University du Quebec a Chicoutimi for identification 
of the ants. We also thank D. Flynn for identification of the Membracidae and Dr. W. 
H. Wagner, Jr., University of Michigan for positive identification of plants. 


LITERATURE CITED 


CLARK, A. H. 1932. The butterflies of the District of Columbia and vicinity. U.S. Natl. 
Mus. Bull. 157, 337 pp. 

CLARK, G. C. & C. G. C. Dickson. 1971. Life histories of the South African lycaenid 
butterflies. Purnell & Sons, LTD, Capetown, 272 pp. 

Common, I. F. B. & F. F. WATERHOUSE. 1972. Butterflies of Australia. Angus & 
Robertson, Sydney, 498 pp. 

Comstock, W. P. 1940. Butterflies of New Jersey. J. N.Y. Entomol. Soc. 48:47-84. 

Downey, J.C. 1961. Myrmecophily in the Lycaenidae (Lepidoptera). Proc. N. Central 
Branch Entomol. Soc. Amer. 16:14—-15. 

1962. Myrmecophily in Plebejus (Icaricia) icarioides (Lepid: Lycaenidae). 

Entomol. News 73:57-66. 

1966. Sound production in pupae of Lycaenidae. J. Lepid. Soc. 20:129-155. 








VOLUME 88, NUMBER 2 133 


HaRVEY, D. J. 1980. Ants associated with Harkenclenus titus, Glaucopsyche lygdamus, 
and Celastrina argiolus (Lycaenidae). J. Lepid. Soc. 34:371-372. 

Ma.icky, H. 1970. New aspects on the association between lycaenid larvae (Lycaeni- 
dae) and ants (Formicidae; Hymenoptera). J. Lepid. Soc. 24:190-202. 

MascHwitz, U., M. Wust & K. SCHURIAN. 1975. Blues’ larvae as sugar suppliers for 
ants. Oecologia 18:17-21. 

McEvoy, P. B. 1979. Advantages and disadvantages to group living in treehoppers 
(Homoptera: Membracidae). Misc. Publ. Entomol. Soc. Amer. 11:1-18. 

Moore, S. 1960. A revised annotated list of the butterflies of Michigan. Occas. Papers 
Mus. Zool., Univ. Mich. 617, 37 pp. 

NEWCOMER, E. T. 1911. The life histories of two lycaenid butterflies. Can. Entomol. 
43:83-88. 

PIERCE, N. E. & P. S. MEAD. 1981. Parasitoids as selective agents in the symbiosis 
between lycaenid butterfly larvae and ants. Science 211:1185-1187. 

Ross, G. N. 1964. Life history studies of Mexican butterflies. III. Early stages of Anatole 
rossi, a new myrmecophilous metalmark. J. Res. Lepid. 3:81—94. 

1966. Life-history studies on the Mexican butterflies. IV. The ecology and 
ethology of Anatole rossi, a myrmecophilous metalmark (Lepidoptera: Riodinidae). 
Ann. Entomol. Soc. Amer. 59:985-1004. 

SCHNEIDER, I. 1960. Soil survey-Montcalm County, Michigan. USDA, SCS, 40 pp. 60 
maps. 

SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with 
special reference to New England. Scudder, Cambridge, 1958 pp. 

Woop, T. K. 1977. Role of parent females and attendant ants in the maturation of the 
treehopper, Entylia bactriana (Homoptera: Membracidae). Sociobiology 2:257-272. 

1982a. Selective factors associated with the evolution of membracid sociality. 

Pp. 175-179, in Breed et al. (eds.). The Biology of Social Insects. Proc. IX Congr. 

International Union for Study of Social Insects, Boulder. 

1982b. Ant-attended nymphal aggregations in the Enchenopa binota complex 

(Homoptera: Membracidae). Ann. Entomol. Soc. Amer. 75:649-653. 











Journal of the Lepidopterists’ Society 
88(2), 1984, 184-137 


THE LIFE HISTORY AND IMMATURE STAGES OF 
AGAPEMA HOMOGENA (SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge #141G, Houston, Texas 77054 


AND 


MICHAEL J. SMITH 
3135 S. Magda, Tucson, Arizona 85730 


ABSTRACT. Agapema homogena is a nocturnal, montane species of saturniid. The 
larvae of homogena feed on Rhamnus californica in Arizona, and have four instars. The 
immature stages are black and yellow with numerous white setae. Adults fly from late 
May to late July in the United States but have been taken as late as mid-September in 
Mexico. The ova are deposited in clusters, and upon hatching the larvae feed gregari- 
ously. In Arizona pupation occurs from September to November. 


Agapema homogena Dyar is a gray to black saturniid of moderate 
size that occurs in Mexico, Arizona, Colorado, New Mexico, and west- 
ern Texas. The species occurs in montane habitats at elevations from 
1500 to 38500 m and in Arizona is most frequently associated with 
mixed oak woodlands (Fig. 1). In this paper the biology of A. homo- 
gena is discussed and the immature stages described for the first time. 
Most of our observations are based on a population in the Santa Cat- 
alina Mts. north of Tucson, Pima Co., Arizona. 


Description of Larvae 
(Figs. 2—4) 


First instar. Head: Black with short white setae; diameter 0.6 mm. Body: Length 5.2 
mm, width 1.8 mm. Ground color black. White setae extend from black scoli. True legs, 
prolegs and spiracles black. 

Second instar. Head: Black with short white setae; diameter 1.4 to 1.5 mm. Body: 
Length 11 mm, width 3 mm. Ground color black and yellow. Dorsal, ventral, and 
intersegmental areas black. Segmental area from dorsal scoli to just ventral of lateral scoli 
yellow with 8 black lines. Black “j” 
dorsally to meet second black line that extends ventrally from mid-dorsal area. 3rd black 
line extends from dorsal area anterior of dorsal scoli, towards dorsolateral scoli but ter- 
minates dorsoanteriorly of dorsolateral scoli. All scoli black and compressed with few 
short black spines and white setae extending from each. White setae extending from 
dorsal scoli of thoracic segments, abdominal segment VIII, and caudal scoli elongated, 
measuring 2 mm or more in length. Prolegs, true legs, and spiracles black. 

Third instar. Head: Black and covered with short white setae; diameter 3.2 mm. Body: 
Length 34-38 mm, width 6 mm. Ground color black and yellow. Ventral and interseg- 
mental area black. Yellow subspiracular line encompasses black lateral scoli and extends 
entire length of larva. Lateral segmental area with 4 vertical yellow stripes. Posterior- 
most stripe extends from subspiracular line dorsally just past dorsolateral scoli then folds 
ventrally to form inverted “v” (often broken) that terminates at spiracle. Line 2 extends 
from spiracle to mid-dorsal area and encompasses dorsolateral and dorsal scoli. On tho- 
racic segments and abdominal segments I, VIII, and IX this line terminates at dorsal 
scoli, therefore, mid-dorsal area of those segments black rather than yellow. Line 3 


VOLUME 38, NUMBER 2 135 





Fics. 1-6. 1, Habitat of Agapema homogena in the Santa Catalina Mts., Pima Co., 
Arizona. Prominent vegetation includes Arctostaphylos sp., Fraxinus sp., Juglans sp., 
Pinus ponderosa, Prunus demissa, Quercus arizonica, Q. emoryi, Q. gambelli, Rhamnus 
californica and Rhus trilobata; 2, lateral view of mature larva; 3, dorsal view of mature 
larva; 4, second instar larvae feeding on Rhamnus; 5, wild cluster of eggs from which 
the larvae have already hatched; 6, two cocoons found in narrow space between rocks. 


anterior to line 2 and extends from posterior edge of spiracle to midway between dor- 
solateral and dorsal scoli. Fourth line anterior to 3rd and extends dorsally from yellow 
subspiracular line to point even with dorsal scoli. Tufts of white secondary setae cover 
mid-dorsal segmental area. All scoli are black and compressed with white setae and few 
black spines extending from each. Dorsolateral and dorsal scoli each with 1, occasionally 
2, setae 5 mm or longer extending from each. Prolegs, true legs, spiracles, and planta 


black. 


136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fourth instar. Head: Black and covered with short white setae; diameter 4.2-5.4 mm. 
Body: Length 58-65 mm, width 11 mm. Ground color black and yellow. Yellow subspi- 
racular line encompasses lateral scoli and extends length of larva. Segmental area with 4 
vertical yellow stripes on each segment as in 3rd instar, but with one exception: the 2nd 
yellow line extends from spiracle dorsally through dorsolateral scoli and terminates at 
dorsal scoli instead of crossing over mid-dorsal area. Mid-dorsal area black with segmental 
tufts of white secondary setae. Lateral scoli appear as yellow verrucae with dense clusters 
of elongated white setae. Dorsal scoli and dorsolateral scoli are distinct but flattened with 
short black spines and 1 to 8 elongated and numerous short white setae. Small clusters 
of white secondary setae extend from white or yellow patch on upper lateral surface of 
prolegs. Prolegs, true legs, and spiracles black. Planta red. 


Discussion 


In southern Arizona the flight season of homogena extends from late 
May to late July. Records from Colorado and New Mexico indicate a 
slightly shorter flight period, while specimens from central and north- 
ern Mexico have been taken through late September (5 km S Temoris, 
Chih. VII-16, VIII-28, [X-19-69). Based on limited material available 
for examination, Ferguson (1972) noted that Arizona specimens are 
smaller and their wing veins more prominent than those from Texas. 
The Arizona specimens that he illustrated are smaller than usual, and 
in general the wing span of material from the two states is similar. The 
difference in wing vein prominence is probably an artifact of reared 
vs. wild specimens since scales rub off the forewing veins of an active 
moth. Material from Colorado to central Mexico has been examined, 
and no consistent geographical trends were found. 

Emergence from the cocoon occurs in the morning. Females begin 
emitting pheromone between 2100 and 2300 h. The pair remain to- 
gether for about an hour, after which the female begins her oviposition 
flight. The ivory eggs are oblong, measure 1 X 2 mm, and are depos- 
ited in clusters near the apical growth (Fig. 5). In the Santa Catalina 
Mts. the larval hostplant is Rhamnus californica ursina (Greene), but 
in Colorado Don Bowman (pers. comm.) has recently collected larvae 
near Steamboat Springs (Routt Co.) on willow. Each egg cluster con- 
tains 45 to 160 eggs. This suggests that females may deposit all of their 
ova in 1, 2 or possibly 3 clusters with 1 or 2 clusters per female being 
the norm. Upon emergence the larvae are gregarious and begin feeding 
on the leaves adjacent to the egg cluster. First instar larvae are black 
but from the second through last instar they are black and yellow with 
long white setae extending from the scoli (Figs. 2-4). Larvae lose most 
of their gregarious tendencies in the last instar. During late August 
larvae in the second through early last instar can be field collected. 
There are four larval instars, and the mature larva measure 55 to 65 
mm. Mature larvae leave the hostplant and wander prior to pupating 
in cracks or crevices among rocks, tree trunks, or man-made structures. 


VOLUME 38, NUMBER 2 137 


Pupation occurs from September to November. Both Mike Collins and 
Mike Van Buskirk (pers. comm.) have observed that field collected 
cocoons, which still contain prepupae or newly molted pupae in No- 
vember or December, were usually parasitized by tachinid flies. Newly 
spun cocoons are light beige but turn a uniform brown with age; the 
change may be hastened by moisture. The cocoon (Fig. 6) is loosely 
woven but somewhat compact. This is in sharp contrast to that of A. 
galbina anona (Ottolengui), which forms a pale brown, bulbous, loose 
mesh cocoon on its hostplant. 

There are many differences between the immature stages of Aga- 
pema and west coast species of Saturnia. Morphologically, the scoli of 
homogena and galbina are reduced and bear far fewer and shorter 
black spines when compared to Saturnia. Agapema larvae are long 
and thin with numerous white secondary setae, especially on the dorsal 
area. The larvae of Saturnia are more compact, and although second- 
ary setae are present, they are inconspicuous. The spines of Saturnia 
are urticating, whereas, those of Agapema are not. In addition Aga- 
pema larvae tend to be gregarious, while Saturnia larvae feed singly 
and are cryptic. The evolution of the New World Saturnia and Aga- 
pema, and their hostplant relationships were recently discussed by 
Tuskes and Collins (1981). In general the larvae of Agapema, especially 
those of homogena, are more divergent from those of Saturnia than 
might be expected considering adult characters and their earlier con- 
generic status (Michener, 1952). 


ACKNOWLEDGMENTS 


We would like to thank Mike Collins for reading the manuscript, Mike Van Buskirk 
who found the colony in the Santa Catalina Mts., and the reviewers for their comments. 


LITERATURE CITED 


FERGUSON, D. C. 1972. The Moths of America North of Mexico. Fasc. 20.2B, Bom- 
bycoidae (in part). E. W. Classey Ltd., London, pp. 155-269, pls. 12-22. 

MICHENER, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemisphere. 
Bull. Amer. Mus. Nat. Hist. 98:339-501. 

TuskEs, P. M. & M. M. CoLLins. 1981. Hybridization of Saturnia mendocino and S. 
walterorum, and phylogenetic notes on Saturnia and Agapema (Saturniidae). J. 
Lepid. Soc. 35(1):1-21. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 138 


GENERAL NOTES 
HOST RECORDS FOR PARATRYTONE MELANE (EDWARDS) (HESPERIIDAE) 


Paratrytone melane (Edwards) is one of several species of Hesperiidae that has become 
increasingly urbanized throughout southern California. According to Thorne (1968, J. 
Res. Lepid. 2(2):148-149), P. melane was not recorded from San Diego County prior to 
1941. In that year, it was first encountered near El Cajon and has since become one of 
our most common urban skippers. In San Diego County, P. melane is generally the first 
skipper on the wing in our residential areas, appearing as early as February. Capture 
records do not indicate a clear broodedness, but rather several or continuous over-lapping 
generations each year, with spring, mid-summer, and fall peaks. 

In mid-March 1982, a female P. melane was observed ovipositing, in what seemed to 
be an indiscriminate manner, on several species of grass, both weedy and lawn, in 
southern San Diego city. A single larva was subsequently reared from an egg deposited 
on goldentop, Lamarkia aurea Linnaeus (Moench) (Poaceae), a common weedy intro- 
duced species. Development was normal, and the adult emerged following a three week 
pupal period. “Indiscriminate’’ oviposition was also observed by Comstock and Dammers 
(1931, Bull. So. Calif. Acad. Sci. 30(1):20-22), but the behavior was that of confined 
females. 

A last instar larva of P. melane was collected and subsequently reared on Saint Au- 
gustine grass, Stenotaphrum secundatum Kuntze (Poaceae), from a lawn in Encinitas, 
California (David Faulkner, San Diego Natural History Museum). Noel MacFarland 
(pers. comm.) also reported rearing P. melane on S. secundatum from the lawn at his 
previous residence in the Santa Monica Mountains, Los Angeles County. 

Larvae of P. melane are known to feed on Bermuda grass, Cynodon dactylon (L.) 
Persoon (Poaceae), in the laboratory (Comstock & Dammers, op. cit.), and observations 
by William McGuire (pers. comm.) of oviposition by P. melane on C. dactylon in 
residential Del Mar, California, suggests its widespread use as a larval host in urban areas. 

Native hosts of P. melane are poorly known; the single report of oviposition on Des- 
champsia caespitosa (L.) Beauvois (Poaceae) (Emmel & Emmel, 1973, Nat. Hist. Mus. 
Los Angeles Co., Sci. Ser. 26:80) is the only record of which I am aware. On one occasion 
the author collected a single last instar larva of P. melane on Carex spissa Bailey (Cy- 
peraceae). Subsequently, Guy Bruyea of Poway, California, also collected and reared a 
single P. melane larva on C. spissa. This native sedge occurs in moist areas generally 
away from the coast and is probably widely used by P. melane in these habitats. 

The presently documented larval hosts of P. melane are restricted to two monocot 
families: Poaceae and Cyperaceae. Several species of grass, both native and introduced, 
are utilized as hostplants or as oviposition substrates. The single known cyperaceous host, 
C. spissa, has exceedingly coarse leaf blades and in this respect seems to indicate that a 
wide range of leaf textures is tolerated by the larvae of P. melane. The ability of P. 
melane to utilize a large number of introduced species has given it the capacity to greatly 
expand its range throughout the urban areas of southern California. 


JOHN W. Brown, Entomology Department, San Diego Natural History Museum, 
P.O. Box 1390, San Diego, California 92112. 


VOLUME 38, NUMBER 2 139 


Journal of the Lepidopterists’ Society 
38(2), 1984, 139 


THE SPHINGID FRENULUM AS A PREDATOR DEFENSE 


Sphingidae, because of their large body size, must present a tempting target to ver- 
tebrate predators. It has been pointed out how tibial spurs can be used to discourage 
would-be predators (Allen, 1982, J. Lepid. Soc. 36:155-157), and in this note I suggest 
an additional defense mechanism. : 

As with Dr. Allen, my knowledge of this mechanism came through personal contact. 
In December 1977, I spent three weeks collecting insects in a remote area of western 
Panama (IRHE camp at Fortuna, Chiriqui Province). Here moths came to light in 
abundance, and the largest were several species of Sphingidae. Since I did not have 
killing jars large enough to hold big moths, my collecting method was to grasp these 
moths by the thorax below the wings and quickly inject several drops of alcohol with a 
hypodermic needle. 

When I collected the largest sphingids (Coctyius and Eumorpha) in this manner, my 
fingers were pricked on several occasions by something extremely sharp. On close ex- 
amination I found that this was caused by the moth’s frenulum. Whenever I grasped the 
moth directly over the wing bases, my fingers would push the forewings up enough to 
expose the frenulum, and at this point it was perfectly positioned to stab into the tips of 
my thumb and forefinger. In the case of the Coctyius and the Eumorpha species at 
Fortuna, the frenulum was thick and stiff enough to pierce my skin. 

The defensive use of the frenulum is, of course, secondary and probably unintentional. 
Nevertheless, my experience leads me to believe that, at least occasionally, sphinx moths 
may be able to escape predators when a well placed jab occurs. The frenulum defense’ 
would be most effective if a bat, toad or lizard were to seize the moth from the front or 
from above. Holding the moth by the front of the thorax would leave the predator out 
of range of the tibial spurs but the struggling moth might be able to stick the frenulum 
into the lining of the predator’s mouth. 


R. W. FLOWERS, Entomology’& Structural Pest Control, Florida A&M University, 
Tallahassee, Florida 32307. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 189-141 


ANOTHER LOOK AT SNOUT BUTTERFLIES 
(LIBYTHEIDAE: LIBYTHEANA) 


The two species of snout butterflies of the southwestern United States and Mexico, 
Libytheana bachmanii (Kirtland) and L. carinenta (Cramer) are commonly confused in 
spite of treatments by Field (1938, J. Kansas Entomol. Soc. 11:124-183), Michener (1943, 
Amer. Mus. Novitates No. 1232), Ehrlich and Ehrlich (1961, How to know the butterflies, 
Wm. C. Brown Co. Publ., Dubuque, Iowa, pp. 174-175), and Heitzman and Heitzman 
(1972, J. Res. Lepid. 10:284—286). They are easily separated in males by the shape of the 
eighth abdominal tergite and less easily (especially in females) by the shape and color- 
ation of the wings. Since the adults have been adequately figured, this note serves to 
illustrate differences in the male eighth abdominal tergites. 

Michener figured the eighth abdominal tergite of L. bachmanii in dorsal and lateral 
views but did not provide a figure of L. carinenta for comparison. As can be seen in 
Figs. 1—4 the species differ in the lateral width of the median apical process and number 
of setae, but more strikingly, in the number of terminal spines. L. bachmanii was found 
to have between 2 and 4 spines (n = 26, mode of 2), while L. carinenta has between 6 


140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-4. 


(many scales have been removed and a few setae broken during preparation); 3 & 4, 
similar view and preparation of L. bachmanii (the crease along the median apical process 
is an artifact from papering the specimen after capture). 


1 & 2, Dorsal view of eighth abdominal tergite of male L. carinenta 


and 9 spines (n = 9, mode of 7). The dorsally projecting spines can be seen by using a 
hand lens or microscope once the overhanging scales have been brushed aside, without 
having to do any dissecting. 

During the morphological investigation of specimens at hand, genitalic dissections were 
done which revealed consistent differences between the species for both sexes. These will 
not be reported here owing to the limited number of specimens investigated and must 
await a comprehensive treatment. However, this look at the terminalia allowed the 
assignment of all but a few female specimens to one species in preference to the other. 


VOLUME 38, NUMBER 2 14] 


Geographically, L. carinenta must be considered a rare find in the United States and 
is not commonly encountered until well below the Tropic of Cancer. L. bachmanii 
broadly overlaps its distribution along the western side of the Gulf of Mexico and is 
found as far south as the Rio Tehuantepec in Oaxaca, Mexico. 

One can only guess as to the function of the terminal spines of males of these butterflies. 
Detailed observations of the mating behavior of snout butterflies might provide the 
answer. Comparative studies of other members of the genus and family of both mor- 
phology and behavior need to be done as part of a revision of this interesting group. 

I would like to thank Drs. H. R. Burke and J. C. Schaffner, Department of Entomology, 
Texas A&M University, for making field studies in Mexico possible. J. Ehrman of the 
Electron Microscopy Center at the University is gratefully acknowledged for his SEM 
work and photography. L. G. Friedlander reviewed the manuscript. 


TIMOTHY P. FRIEDLANDER, Department of Entomology, Texas AUM University, 
College Station, Texas 77843-2475. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 141-142 


COMMUNAL ROOST FIDELITY IN HELICONIUS CHARITONIA: 
COMMENTS ON A PAPER BY 
DRS. D. A. WALLER AND L. E. GILBERT 


In the recent paper by Waller and Gilbert appearing on the pages of this journal (J. 
Lepid. Soc. 36:178-184), the authors failed to include other substantial data sets on 
communal roosting in Heliconius charitonia and related aspects of this butterfly’s pop- 
ulation biology which have significant bearing on their conclusions and comments (Young 
& Thomason, 1975, J. Lepid. Soc. 29:243-255; Cook, Thomason & Young, 1976, J. Anim. 
Ecol. 45:851-868). 

Waller and Gilbert imply that at least a portion of the daily instability in roost mem- 
bership observed for two other studies of H. charitonia in Costa Rica (Young & Carolan, 
1976, J. Kansas Entomol. Soc. 49:346-359; Young, 1978, Entomol. News 89:235-243) was 
due to disturbance of butterflies for marking, something they apparently avoided in their 
study. This is a serious accusation, one that is not merited as seen by the examination of 
Young and Thomason (op. cit.) and Cook et al. (op. cit.), two additional Costa Rican 
studies of the same organism not cited by Waller and Gilbert, and ones that report a 
significant amount of both population cohesiveness and fidelity to communal roosts. 

There is no doubt that butterflies are disturbed to some extent by the handling effects 
associated with marking, a condition that I seriously doubt even Waller and Gilbert could 
have avoided entirely in their study. The same techniques associated with marking, 
however, were used in all of the Costa Rican studies cited above, and therefore, any 
handling effects causing roost disturbance would have been the same for all data sets. 
Yet Young and Thomason (op. cit.) reported for Roost A in that study, that of 69 but- 
terflies marked, 36 were seen again at least once, and 23 seen from one to three times 
on subsequent days of observation. We concluded that roost fidelity can be high in H. 
charitonia, but that the spatial distribution of multiple roosts within the same home 
range area used by the butterflies on any one roost results in considerabe “exchanges” 
among roosts on a day-to-day basis. Admittedly, this level of roost fidelity is still somewhat 
lower than the findings of Waller and Gilbert in Mexico, yet higher than observed for 
other roosts in Costa Rica (Young & Carolan, op. cit.). Furthermore, the study of Cook 
et al. (op. cit.) on H. charitonia population dynamics spanned a period of 155 days and 
involved the marking of 586 butterflies and concluded that the movement of individual 


142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


butterflies is regulated largely by the locations of communal roosts and adult and larval 
food resources. That study also revealed a fractionation of the population into several 
subpopulations but with considerable interchanges of marked butterflies between areas 
of habitat occupied by different subpopulations. The obvious inference from such results 
is the shifting dependency of individual butterflies among several communal roost sites 
within a relatively small area of habitat. Waller and Gilbert (op. cit.) did not mention 
the occurrence of other roosts within the vicinity of those adult pollen-source plants 
visited principally by unmarked individuals of H. charitonia. Given the results of Cook 
et al. (op. cit.), other roosts most likely existed in the generai vicinity of the home range 
area occupied by these unmarked butterflies. 

The results of Young and Thomason (op. cit.) indicated that there can sometimes occur 
considerable individual variation in the tenacity of H. charitonia to a particular roost 
site. Genotypic differences among individual butterflies may ultimately explain such 
patterns (Young and Thomason, op. cit.). In the absence of such data, however, it is safe 
to conclude tentatively that in some tropical regions occupied by H. charitonia, the 
degree of fidelity to a particular roost site is highly dependent upon (1) the availability 
of multiple roosts within the area, (2) the positioning of different home ranges occupied 
by different subpopulations relative to one another, and (3) the abundance and spatial 
distribution of adult and larval food resources within home range areas. Given the find- 
ings of Young and Thomason (op. cit.) and Cook et al. (op. cit.), I believe that it is 
erroneous on the part of Waller and Gilbert (op. cit.) to suggest that the patterns of roost 
instability reported in Young and Carolan (op. cit.) and Young (op. cit.) as being due to 
disturbance incurred while marking butterflies. Waller and Gilbert did not discuss the 
results of Young and Thomason (op. cit.) relative to their interesting data. Had they done 
so, they might have been able to suggest that the observed high fidelity of butterflies to 
the single roost they studied was possibly due to the absence of a second roost within the 
same home range or at the periphery of a contiguous home range associated with the 
unmarked butterflies they saw at patches of adult pollen-sources far removed from the 
vicinity of the roost in question (a projected spatial arrangement of home ranges and 
roosts that would probabiy preclude frequent exchanges of marked butterflies among 
different roosts). In doing so, they would have justifiably assigned an equal weight or 
error factor to disturbance of butterflies during marking in both their study and the Costa 
Rican studies discussed here. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 142-143 


RAINSTORM BEHAVIOR OF PIPEVINE SWALLOWTAILS, 
BATTUS PHILENOR (L.) 


While collecting near Laredo, Texas in mid-afternoon, 12 June 1981, we took shelter 
in our car in advance of a rainstorm approaching from the southeast. The car was parked 
among mesquite trees, Prosopis glandulosa Torr., and we watched as six pipevine swal- 
lowtails, Battus philenor (L.), buffeted by a brisk wind, came together in a little group 
on one of the trees from the otherwise sparse population of this butterfly in the area. 
With the sun in the opposite direction from the storm, no darkening of skies had occurred 
at the time the assembly was initiated. Individuals were all about 12 feet from the ground, 


VOLUME 38, NUMBER 2 143 


separated from each other by inches to a foot or two. All located themselves on the lee 
side of twigs, head upward and wings folded together over their backs. After the heavy 
rain shower they gradually disassembled, fanning their wings before flying away one- 
by-one. One individual moved for a time to another tree and repositioned itself on a 
twig but on the side of the continuing southeast breeze, with wings spread apart and not 
fanning. 

In their paper on roost recruitment and resource utilization by Heliconius charitonia 
L. near Vera Cruz, Mexico, D. A. Waller and L. E. Gilbert (1982, J. Lepid Soc. 36:178- 
184) review hypotheses on communal! roosting and comment that Heliconius roosting 
behavior is one of the major remaining mysteries of lepidopteran biology. In relation to 
our observations, Gilbert (pers. comm.) mentions that the roosts at Vera Cruz, where 
daily rains were the rule, formed earlier when storms occurred in the early afternoon. 
He has also seen such roosting in B. philenor and Danaus gilippus (Cramer) around 
Catarina, Dimmit Co., Texas. 

Our observations were made during a one-day trip and without opportunity for more 
extended observation. While difficult in south Texas because of sporadic rainfall, further 
observation of roosting behavior on days with and without afternoon thunderstorms will 
be necessary to extend and explain our observations for Battus and other species. It 
would be interesting to know whether the butterflies we observed returned to the same 
place for roosting at night. 


JAMES E. GILLASPY AND JOHNNY R. Lara, Department of Biology, Texas A&I Uni- 
versity, Kingsville, Texas 78363. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 143-144 


WESTERN RANGE EXTENSIONS FOR ANISOTA CONSULARIS 
(SATURNIIDAE) REPRESENTING NEW STATE RECORDS 
IN MISSISSIPPI AND LOUISIANA 


Until recently, the known distribution of Anisota consularis Dyar was limited to a 
few scattered records from Florida. The inability of reviewers to correctly separate A. 
consularis from its Floridian congeners only further limited our knowledge of the species’ 
range. Kimball (1965, Lepidoptera of Florida, p. 69) readily admitted the limitations of 
his knowledge of A. consularis and Ferguson (1971, Moths of North America, Fascicle 
20(2A), Bombycoidea: Saturniidae (Part), pp. 63-84) had difficulty distinguishing be- 
tween A. consularis and Anisota stigma Fabricius. 

The revision of the genus by Riotte and Peigler (1980(81), J. Res. Lepid. 19(3):101- 
180) offers the first taxonomic understanding of A. consularis and corrects many of the 
previously published mis-identifications. In addition, they offer records of A. consularis 
from Long and Bulloch counties of coastal Georgia. These captures are the only previ- 
ously published reports of A. consularis occurring outside of Florida. 

Several years ago, through the generosity of curator Patricia Ramey, the author ex- 
amined the Anisota in the Mississippi Entomological Museum at Mississippi State Uni- 
versity. A previously undetermined female collected by C. C. Greer at Gulfport, Harrison 
County, Mississippi, on 1 September 1916, was identified by the author as A. consularis. 
This specimen represents a new state record and westward range extension for A. con- 
sularis. 

Recently, the author also examined the Anisota in the private collection of Vernon A. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brou, Edgard, Louisiana. Among the material was a female A. consularis collected by 
Brou on 3 August 1978, at Fluker, Tangipahoa Parish, Louisiana. This capture also 
constitutes a state record and further extends the western range of A. consularis. 

It is noteworthy that both collecting locales are from coastal areas. Additional captures 
of A. consularis should be anticipated in those areas of Louisiana, Mississippi, and Ala- 
bama, where a mild climate is maintained by the warming influence of the Gulf of 
Mexico. 


JIM TUTTLE, 728 Coachman #4, Troy, Michigan 48083. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 144-146 


A BILATERAL SEXUAL MOSAIC OF MITOURA GRYNEUS (LYCAENIDAE) 


A bilateral sexual mosaic of the Olive Hairstreak, Mitoura gryneus (Hiibner) was 
collected on 1 August 1981, from Red Cedar, Juniperus virginia, near Lynx, Adams 
County, Ohio. The only other specimen collected on that date was a typical female. 

The right half of the specimen is male in appearance and is strongly suffused with 
gold scales (Fig. 1). The left half has a mixture of male and female characters and is 
dark brown with a dusting of orange-brown scales, a female characteristic. The right 
scent pad is oval and measures 1.98 x 0.71 mm (Fig. 2). The left scent pad is sickle- 
shaped and measures 1.54 x 0.39 mm. A small sample (n = 8) of typical scent pads from 
southern Ohio was all oval and averaged 2.01 + 0.113 x 0.73 + 0.026 mm. No differ- 
ences in pattern can be detected on the ventral wing surfaces. 

The genitalia of the mosaic were dissected and compared to typical male genitalia 
from southern Ohio (Fig. 2). The right half appears to be typically male, but the left 
half has several abnormalities. The halves of the uncus are not fused medially, and the 
left half is largely unsclerotized and dorsally enlarged. The left valva is narrowed basally 
but is otherwise well developed. A partially sclerotized projection from the left vinculum 





Fic. 1. Bilateral sexual mosaic of Mitoura gryneus. 


VOLUME 38, NUMBER 2 145 





Fic. 2. Male genitalia and scent pads of Mitoura gryneus: A, sexual mosaic genitalia, 
posterior ventral view; B, typical genitalia, posterior ventral view; C, sexual mosaic 
genitalia, lateral view; D, typical genitalia, lateral view; E, left mosaic scent pad; F, right 
mosaic scent pad. F, falx; P, pedunculus; S$, saccus; U, uncus; V, valva; Vi, vinculum. 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


is similar in structure and shape to the valvae and may represent the development of a 
second left valva. At this projection’s point of attachment the vinculum is very broad 
and flattened. Near the juncture of the vinculum and pedunculus a heavily sclerotized 
rod projects anteriorly. The rod has no apparent counterpart in typical male or female 
genitalia. The aedeagus does not differ from that of typical males. 

I thank Norman Reichenbach and Dr. N. Johnson, Ohio State University, for reviewing 
the manuscript. 


JOHN A. SHUEY, The Ohio State University, Department of Entomology, 1735 Neil 
Avenue, Columbus, Ohio 43210. 


Journal of the Lepidopterists’ Society 
38(2), 1984, 147 


BOOK REVIEWS 


THE LIFE HISTORIES OF BUTTERFLIES OF JAPAN. VOLUME I. PAPILIONIDAE, PIERIDAE, 
DANAIDAE, by Haruo Fukuda, Eiichi Hama, Takeshi Kuzuya, Akira Takahashi, Mayumi 
Takahashi, Ban Tanaka, Hiroshi Tanaka, Mario Wakabayashi and Yasuyuki Watanabe. 
xxii + 277 pp., 64 col. pls., 1982. Hoikusha Publishing Co., Ltd., 17-3, 1-chome, Uemachi, 
Higashi-Ku, Osaka, 540, Japan. (Price not stated in review copy.) 


This book treats the up-to-date knowledge of the early stages, adult behavior and 
distribution of all resident and vagrant species of the families mentioned. The area 
covered ranges from Hokkaido southwest to Iriomote-Jima. Thus, the entire chain, rang- 
ing from Arctic-Alpine to Subtropical conditions, is included. The main text is in Japanese 
and appears to be authoritative as extensive references are cited and topics such as 
changes in distribution and mortality curves are included. 

For the majority of our membership there are 12 pages of capsule species accounts 
with cross reference to the color plates and 16 pages and distribution maps. 

As we have come to expect from Japanese books, the color plates are of high quality 
with the colors appearing sharp and true. For each species the adults as found in nature, 
egg, larva, and pupa are portrayed. Often the habitat or host plant is also included. The 
adults are often shown in natural behavioral activities such as mating, oviposition, taking 
moisture or nectaring. The pictures are excellent but occasionally not clear. 

For those interested in the many parallels in the life history traits between our species 
and those of Japan, this book is a must. 


PAUL A. OPLER, Division of Biological Services, U.S. Fish and Wildlife Service, Wash- 
ington, D.C. 20240. (Current address: U.S. Fish and Wildlife Service, Colorado State 
University, Fort Collins, Colorado 80523.) 


Journal of the Lepidopterists’ Society 
38(2), 1984, 147-148 


THE BUTTERFLIES OF THE YEMEN ARAB REPUBLIC, by Torben B. Larsen. Royal Danish 
Academy of Sciences and Letters, Kobenhavn. Biologiske Skrifter 23(3). 62 pp. 1982. 120 
Danish kroner. 


The Middle East is, sadly, best known around the world for its seemingly endless cycles 
of vengeance and violence. Between the wars, a few hardy souls have been able to do 
pioneering work on the lepidopteran faunistics of the region; foremost among them is 
Torben Larsen. Larsen’s 1974 book Butterflies of Lebanon remains a model of how to 
do a regional fauna correctly. He continued his tradition of excellence with work in 
Oman, east Jordan, and Saudi Arabia, and now with this little monograph of the Yemeni 
fauna. Naive Americans who think the Persian Gulf region consists of bare dunes and 
date-palm oases—and little else—have a lot to learn from this work. Larsen provides fine 
discussions of climate, vegetation, butterfly distributions and seasonality. There are few 
North American faunistic papers that match this for the sophistication of the ecogeo- 
graphic presentation. 

The main body of the text—the species accounts—includes some geographical sur- 
prises. This is hardly surprising. Although there has been an unexpectedly long history 
of collecting in Yemen, the aggregate data are scanty. Larsen spent five weeks in the 
country in prime season and got 101 species. He missed only 5 species recorded by others 
for the country, and added 31 to the list. He was also able to find some new entities: the 
new taxa, for which excellent descriptions and figures are provided, are Neptis serena 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


annah, Cacyreus niebuhri, Lepidochrysops forsskali, and the genus Tuxentius, removed 
from Castalius. There is also a description of a unique, unnamed female of a new 
Lepidochrysops. All the new taxa and others of particular interest are shown on two 
color plates, which include three surprisingly lush habitat photos. 

There is an appendix by A. H. B. Rydon: “Taxonomic notes on some members of the 
Charaxes viola group, with descriptions of three new species from the Yemen Arab 
Republic and Ethiopia,” with one color and two black-and-white plates. 

When Larsen has completed his series of monographs for the region, biogeographers 
and ecologists will be able to look for general organizational rules for butterfly faunas in 
desert and seasonal-arid climates; already some hints of order are beginning to emerge. 
Now, if only the people of the region could be persuaded to put down their guns and 
go butterfly-hunting instead! 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Date of Issue (Vol. 38, No. 2): 16 August 1984 


EDITORIAL STAFF OF THE JOURNAL 
THoMas D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MacbDa R. Papp, Editorial Assistant 
DouGLas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 
NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
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196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


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CONTENTS 


LIFE HISTORIES OF TAENARIS (NYMPHALIDAE) FROM PAPUA NEW 
GuINEA. Michael Parsons 2 


A NEw SPECIES OF SIMILIPEPSIS AND TAXONOMIC PLACEMENT 
OF THE GENUS (SESIIDAE). Ping Yuan Wang 


NOTES ON THE LARVA OF CARGIDA PYRRHA (NOTODONTIDAE). 
George L, Godfrey ee 


THE LARVA OF AUTOGRAPHA FLAGELLUM (WALKER) (NOCTUI- 
DAE: PLUSIINAE). . Kenneth Neil 2. 


A NEw HAWKMOTH FROM QUINTANA ROO, MExico. Vernon 
Antoine Brou; Jr). re 


NATURAL History NOTES FOR TAYGETIS ANDROMEDA (CRAMER) 
(SATYRIDAE) IN EASTERN Costa Rica. Allen M. Young ... 


THE LIFE-HISTORY OF ACTIAS MAENAS (SATURNIIDAE). Wolf- 
gang A. Ndssig & Richard Steven Peiglher ee 


MYRMECOPHILY IN THE EDWARD’S HAIRSTREAK BUTTERFLY 
SATYRIUM EDWARDSII (LYCAENIDAE). R. P. Webster & M. 
C, Nielsen 220 22 i 


THE LIFE HISTORY AND IMMATURE STAGES OF AGAPEMA 
HOMOGENA (SATURNIIDAE). Paul M. Tuskes & Michael J. 
Sotith 2. 3 OR a SN 2 


GENERAL NOTES 


Host records for Paratrytone melane (Edwards) (Hesperiidae). John W. 
Browrrn i Eh ON 
The sphingid frenulum as a predator defense. R. W. Flowers ccc 
Another look at snout butterflies (Libytheidae: Libytheana). Timothy P. 
Friedlander 
Communal roost fidelity in Heliconius charitonia: comments on a paper by 
Drs. D. A. Waller and L. E. Gilbert. Allen M. Youreg ...ce eee 
Rainstorm behavior of pipevine swallowtails, Battus philenor (L.). James 
E. Gillaspy <> Johnny R, Dare i008 


Western range extensions for Anisota consularis (Saturniidae) representing 


new state records in Mississippi and Louisiana. Jim Tuttle WW... 
A bilateral sexual mosaic of Mitoura gryneus (Lycaenidae). John A. 
Shree yp A a rr 


124 


134 


Volume 38 1984 Number 3 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





24 April 1985 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Don R. Davis, President LEE D. MILLER, 

Vitor O. BECKER, Vice President Immediate Past President 
JAVIER DE LA MAza E., Vice President JULIAN P. DONAHUE, Secretary 
JOHN C. DOWNER, Vice President Eric H. METZLER, Treasurer 


Members at large: 


F. S. CHEw J. M. BuRNsS B. A. DRUMMOND 
G. J. HARJES F. W. PRESTON J. LANE 
E. H. METZLER N. E. STAMP R. K. ROBBINS 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
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itate the exchange of specimens and ideas by both the professional worker and the 
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Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
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Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- 
temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- 
sion in the Sesiidae fascicle for the Moths of America North of Mexico. The dusky 
clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of 
willows and poplars. 


JouRNAL OF 
Tue LeEePIDOPTERISTS’ SOCIETY 


Volume 38 - 1984 Number 3 


Journal of the Lepidopterists’ Society 
38(3), 1984, 149-164 


SOD WEBWORM MOTHS (PYRALIDAE: CRAMBINAE) 
IN SOUTH DAKOTA 


B. MCDANIEL,! G. FAUSKE! AND R. D. GUSTIN? 


ABSTRACT. Twenty-seven species of the subfamily Crambinae known as sod web- 
worm moths were collected from South Dakota. A key to species has been included as 
well as their distribution patterns in South Dakota. 


This study began after damage to rangeland in several South Dakota 
counties in the years 1974 and 1975. Damage was reported from Cor- 
son, Dewey, Harding, Haakon, Meade, Perkins, Stanley and Ziebach 
counties. An effort was made to determine the species of Crambinae 
present in South Dakota and their distribution. Included are a key for 
species identification and a list of species with their flight periods and 
collection sites. 


MATERIALS AND METHODS 


Black light traps using the General Electric Fluorescent F,; T8 B1 15 
watt bulb were set up in Brookings, Jackson, Lawrence, Minnehaha, 
Pennington and Spink counties. In Minnehaha County collecting was 
carried out with a General Electric 200 watt soft-glow bulb. Daytime 
collecting was used in several localities. Material in the South Dakota 
State University Collection was also utilized. For each species a map 
is included showing collection localities by county. On the maps the 
following symbols are used: @ = collected by sweepnet. © = collected 
by light trap. 


Key to South Dakota Crambinae 


la. R, stalked 
Fb. RK; arising directly from discal cell 22 


1 Plant Science (Entomology) Department, South Dakota State University, Brookings, South Dakota 57007. 
* Northern Grain Insect Research Laboratory, Brookings, South Dakota 57006. 


150 


2a. 
2b. 
oa. 
3b. 
Aa. 
Ab. 
oa. 


Ob. 


6a. 
6b. 


7a. 
Tb. 
Sa. 
8b. 
Qa. 
Qb. 


10a. 


10b. 


lla. 
1lb. 


Za 
12b. 
13a. 


13b. 
14a. 
14b. 
15a. 
15b. 
16a. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Male antennae simple, serrate, lamellate 0 3 
Male antennae pectinate 19 
Forewing with discal area silvery white 00 4 
Discal area not silvery white ..... eee 12 
Forewing silvery white, no stripe (EEE 5 
Forewing with silvery white stripe or stripes 0 6 
Dark scaling in median area, complete row of terminal black 

SOUS Sen bts eee ns eee Microcrambus elegans Clemens 
Dark scaling absent, immaculate silvery white 00. 

7 ann see ne Reon Wes i a Crambus perlellus innotatellus (Walker) 
Forewing with single silvery stripe 00 t 
Forewing with one silvery stripe in the discal cell, one silvery 

stripe along costa uw. 1] 
A white patch beyond discal silvery stripe 0 8 
Without white patch 000. eee 9 
Stripe extends beyond costal inception of subterminal line; 

forewing apex falcate Crambus pascuellus floridus Zeller 
Stripe not reaching costal inception of subterminal line; apex 

SOMLALC sulin, S Wee Miho aaa t he 2 Le Crambus alboclavellus Zeller 
Wing base with stripe and brown area above discal stripe 

nearly equal in width Crambus praefectellus Zincken 
Wing base with costal brown area reduced to a narrow line 


Gnathos narrow at base, broadening distally, appearing spoon 
shaped; subterminal area of wing as dark as median area, 
usually with four black dashes ......... Crambus ainslieellus Klots 

Gnathos narrow throughout; subterminal area paler than me- 
dian area, usually with five black dashes 0 
AI radar hk SRE Nt SR Crambus leachellus Zincken 

Terminal line preceded by black dots 
et SN Baad asl dren ect Crambus agitatellus Clemens 

Terminal line preceded by black dashes 
se ac Nei at 2 Vc Crambus laqueatellus Clemens 

Fringe metallie gold 2. 13 

Fringe not metallic gold 2. eee 16 

Terminal line replaced on lower half of wing by dots ............. 

Sale MG SA UL EAN ates NP cae a Chrysoteuchia topiaria (Zeller) 
Terminal row of dots:complete |. | ee 14 
White scaling along cubitus Crambus coloradellus Fernald 
White scaling along cubitus absent 15 
Front: conic¢ah 5 Ae Agriphila vulgivagella (Clemens) 
Front flattened, -.22 = -..a ean a Agriphila ruricolella (Zeller) 
Lower half of forewing along inner margin darker than discal 

area 


VOLUME 38, NUMBER 38 [St 


16b. Lower half of inner margin of forewing not darker than 
CGC AARC A Mtl: wwe os ya Goes Pediasia luteolella (Clemens) 


Three forms are recorded from South Dakota and are separated as follows: 


Horewine light yellow-brown 2 P. I. luteolella (Clemens) 
IRerewine dark brows 80.8). Sk APs POA P. I. caliginosella (Clemens) 
OFEWING PTAY 2 dBc raigk We Dalits Eee SIL eee P. |. zeella (Fernald) 


17a. Fringe cut by white opposite veins ... Pediasia trisecta (Walker) 
17b. Fringe ground color, not cut by white opposite veins _.. 18 
18a. Three terminal black dots; inner margin sprinkled with black 
SEES te res SUE Be Pediasia dorsipunctella (Kearfott) 
18b. Seven terminal black dots Pediasia mutabilis (Clemens) 
19a. Male antennae bipectinate Thaumatopsis pexellus (Zeller) 
Ripeeviale antennae unipectinate .22 22 20 
20a. A black stripe from base to apex of forewing 
__ es ee adres Thaumatopsis fernaldellus Kearfott 
PRES kestnioe 2USCNl 6 eh pe ela 21 
HaaCanitus white scaled;:wing brown 2.2 
eee en Thaumatopsis pectinifer Zeller 
21b. Cubitus not set off by white scaling; wing light brown to 
oubibe erates. AIF Meri eer yoy Thaumatopsis repandus Grote 
ee@lecHinalusen(g ee eves it hk ee ee 23 
upE@lecigmReseit se Te 24 
23a. Palpi more than three times as long as head; hindwings white 
Me fo Thopeutis forbesellus (Fernald) 
23b. Palpi less than twice as long as head; hindwings brownish 
Tea eee Occidentalia comptulatalis (Hulst) 
24a. Wings crossed by two yellow stripes 
pram As Peles. Hie) iy ET os Euchromius californicalis (Packard) 
Zane wines without two yellow stripes 9 25 
25a. Forewings with vein 3A present; silvery white with brown 
ae itch oe Sal Og We LOANS 8 2TH Platytes vobisne Dyar 
PPE Orewines with GA aAbsemb cessed Ne 2 26 
26a. Vein R, arising from discal cell; color silvery white 
oe I ARAL 2 ccc aed Nahe eee Argyria nivalis (Drury) 
26b. Vein R, stalked with R;,,; color brown with light colored 
CET] Gee ae Eee Ey Pee Sora Eoreuma crawfordi Klots 


Subfamily Crambinae 


Crambus praefectellus praefectellus Zincken 
(Figs. 1C-2, 4a & b) 
Records. 99 specimens from Brookings, Lawrence, Minnehaha counties. 


Flight period. Bivoltine with peaks in June and August, extreme dates 2 May to 18 
September. 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Remarks. Klots (pers. comm.) calls this subspecies an eastern and Rocky Mountain 
form. He stated that the Rocky Mt. race is larger and brighter. He called attention to 
the wide brown costal border from base of the forewing. 


Crambus leachellus Zincken 
(Figs. 1A-6, 4g & h) 


Records. 56 specimens from Brookings, Minnehaha, Spink counties. 

Flight period. Peaked late in September, extreme dates 31 August to October. 

Remarks. Klots (pers. comm.) stated that this species is almost continent-wide; often 
very common to abundant. He states it is of some economic importance in lawns. More 
or less continually on wing over a long period of time. Klots considers the color and 
pattern usually indistinguishable from C. ainslieellus but genitalia very distinctive. We 
have found the gnathos to be a distinguishing character in separating C. leachellus from 
C. ainslieellus in South Dakota material. 


Crambus ainslieellus Klots 
(Figs. 2A, 4c & d) 


Records. 135 specimens from Brookings, Custer, Dewey, Haakon, Harding, Jackson, 
Lawrence, Meade, Minnehaha, Spink, Ziebach counties. _ 

Flight period. Most records in mid-September, extreme dates 30 August to 24 Septem- 
ber. 

Remarks. Klots (pers. comm.) considers C. ainslieellus like C. leachellus to be almost 
continent-wide in distribution. He states that C. ainslieellus is rare or uncommon. How- 
ever, we have found it to be the more common species in the western section of South 
Dakota. 


Crambus laqueatellus Clemens 
(Figs. 1A-5, 4i & j) 


Records. 3 specimens from Brookings, Minnehaha counties. Dates of capture were 31 
June and 3 August. 

Remarks. Klots (pers. comm.) regards C. laqueatellus as fundamentally an eastern 
species. He states that it is often the first Crambus to fly in late spring. C. laqueatellus 
was only found in the eastern portion of South Dakota and only 3 specimens were 
collected. It is normally associated with lawn grasses and wet meadows. 


Crambus perlellus innotatellus Walker 
(Figs. 2B, 7c) 


Records. 20 specimens from Brookings, Jackson, Lawrence, Meade, Minnehaha, Pen- 
nington, Shannon counties. 


Flight period. Most records in early August, extreme dates 19 July to 11 September. 


Crambus agitatellus Clemens 
(Figs. 1A-2, 5f) 


Records. 2 specimens from Minnehaha County dated 16, 21 June. 

Remarks. Klots (pers. comm.) said C. agitatellus is the conventional name now used 
in literature, but this will be corrected when he publishes his work dealing with the 
Crambinae. 


VOLUME 38, NUMBER 3 153 























Fic. 1. Distribution data for: A) 1—Crambus alboclavellus, 2—Crambus agitatellus, 
3—Crambus pascuellus floridus, 4—Microcrambus elegans, 5—Crambus laqueatellus, 
6—Crambus leachellus, 7—Crambus coloradellus. B) 1—Thaumatopsis pectinifer, 2 
Thaumatopsis repandus, 3—Occidentalia comptulatalis, 4—Thopeutis forbesellus. C) 
1—Argyria nivalis, 2—Crambus praefectellus. 





Crambus alboclavellus Zeller 
(Figs. LA-1, 4e & f) 


Records. A single specimen collected from Minnehaha County dated 15 July. 

Remarks. Klots (pers. comm.) regards this species as often abundant in East, rarer 
westward. He said C. alboclavellus Zeller is the conventional name for this species; 
however, it is incorrect. He regards C. alboclavellus as difficult to distinguish from C. 
agitatellus, but genitalia are distinctive. We have only studied 3 specimens of C. albo- 
clavellus and C. agitatellus; therefore our use of the presence or absence of a white 
patch beyond the single stripe on the forewing may not be a reliable character; genitalia 
were distinctive for the South Dakota material. 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Crambus pascuellus floridus Zeller 
(Figs. 1A-3, 5a) 


Records. 4 specimens from Minnehaha County. 

Flight period. Specimens collected on 2 July and 7 August. 

Remarks. Klots (pers. comm.) said this species exists in the northern % of continent. 
He states that the nominate subspecies is Palaearctic and that C. pascuellus floridus is 
definetely a northern species, but found in the mountains of southern areas. 


Microcrambus elegans (Clemens) 
(Figs. 1A-4, 7g) 


Records. 48 specimens from Minnehaha County. 
Flight period. Most records in mid-July, extreme dates 28 June to 10 September. 


Crambus coloradellus Fernald 
(Fig. 1A-7) 


Records. 2 specimens from Buffalo and Jackson counties dated 7 August. 

Remarks. Klots (pers. comm.) stated that coloradellus is not congeneric with other 
members of the genus Crambus and that he is in the process of erecting a new combi- 
nation for this species. However, for the purpose of this paper it will be treated as a 
member of the genus Crambus. 


Chrysoteuchia topiaria (Zeller) 
(Figs. 2C, 4k & 1) 


Records. 329 specimens were collected from Brookings, Jackson, Lawrence, Minne- 
haha, Pennington counties. 

Flight period. Most records for South Dakota are in mid-July, extreme dates 2 May 
to 25 August. 

Remarks. Klots (pers. comm.) states that this species has, until recently, been regarded 
as a North American race of the European “Crambus”’ hortuellus (Huebner) and is still 
so treated in the literature. It has been recorded as a pest on cranberry; however, it 
occurs widely where there is no cranberry, often it is abundant in grasslands. According 
to Klots there may be two “sibling” species, or “food plant” species. The species is 
northern but exists continent-wide. 


Agriphila vulgivagella (Clemens) 
(Figs. 2D, 5g & h) 


Records. 121 specimens from Brookings, Dewey, Codington, Harding, Jackson, Law- 
rence, Meade, Minnehaha, Spink counties. 

Flight period. Peak in early September, extreme dates 19 July to 18 September. 

Remarks. In the literature this species is recorded as Crambus vulgivagellus the well 
known pest called “Vagabond Crambus.”’ It is most often confused with A. ruricorella 
here in South Dakota. 


Agriphila ruricorella (Zeller) 
(Figs. 2E, 5e) 


Records. 188 specimens from Clay, Dewey, Hamlin, Harding, Jackson, Lawrence, 
Meade, Minnehaha, Pennington, Shannon, Spink counties. 

Flight periods. Most records in August, extreme dates 11 July to 23 September. 

Remarks. A ruricorella is a smaller form than A. vulgivagella and often has some 


VOLUME 38, NUMBER 3 Lap 








Fic. 2. Distribution data for: A—Crambus ainslieellus, B—Crambus perlellus in- 
notatellus, C—Chrysoteuchia topiaria, D—Agriphila vulgivagella, E—Agriphila ruri- 
corella, F—Pediasia luteolella. 


transverse markings on the forewings. The most distinguishing character of A. ruricorella 
is the flattened front. In A. vulgivagella the front is produced and conical. 


Pediasia luteolella (Clemens) 
(Pigs. 2H Sb, c, d, i) 


Records. 47 specimens from Brookings, Hughes, Jackson, Lawrence, Minnehaha, Spink 
counties. 

Flight period. Most records in July, extreme dates 11 June to 31 August. 

Remarks. Klots (pers. comm.) states regarding two specimens sent to him “The speci- 
mens are not typical, but in this mess they are seldom so.”’ He referred to the specimens 
as a continent-wide species complex, or superspecies, that includes P. |. zeella Fernald 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and P. I. caliginosella (Clemens). Color ranges from almost unmarked yellow to dark 
sooty brown, sometimes with contrasting marks, sometimes almost unmarked. Klots re- 
gards some of this group to be of economic importance. Some will damage sprouting 
corn. He expects to be putting this group in a new genus. In this paper we have treated 
zeella (Fig. 5c), and caliginosella (Fig. 5b), as subspecies and devised a key utilizing 
color to separate P. I. luteolella, P. |. caliginosella and P. |. zeella; this has worked well 
for our South Dakota material but may not hold up regarding other regions. 


Pediasia trisecta (Walker) 
(Figs. 3A, 7e & f) 


Records. 3597 specimens from Brookings, Buffalo, Clay, Dewey, Codington, Fall Riv- 
er, Harding, Hyde, Jackson, Jones, Lawrence, Meade, Minnehaha, Pennington, Spink, 
Stanley, Todd counties. 

Flight period. Bivoltine possibly trivoltine with peaks in July and September, extreme 
dates 7 May to 13 October. 

Remarks. This species, until recently, has been known in the literature as Crambus 
trisectus. This is a very common webworm associated with lawns throughout South 
Dakota. It is very abundant during peak periods and can cause economic damage to 
grass lawns. According to Klots (pers. comm.) this species has white veins; often has 
reduced or short white streaks at the margin or in fringe area. A couple of very close 
species with sexual dimorphism exist. Also there is much variation between the 2-3 
generations. This species is a continuous flyer. Klots warns that specimens with no traces 
of the white streaks should be carefully examined. They may be Pediasia laciniella 
(Grote). 


Pediasia dorsipunctella (Kearfott) 
(Figs. 3B, 6e & f) 


Records. 191 specimens from Brookings, Jackson, Lawrence, Pennington, Spink coun- 
ties. 

Flight period. Most records in mid-August, extreme dates 19 July to 18 September. 

Remarks. This species is smaller than P. trisecta, there are no white streaks in fringe 
of forewing. The male genitalia will separate P. dorsipunctella from P. trisecta in that 
the gnathos lack the terminal hook. To separate females the terminal fringe of forewing 
cut with white streaks is the important distinguishing character. 


Pediasia mutabilis (Clemens) 
(Figs. 8C, 6g & h) 


Records. 58 specimens from Brookings, Brown, Duel, Lawrence, Minnehaha, Spink 
counties. 

Flight period. Bivoltine with peaks by early July and early September, extreme dates 
10 June to 18 September. 

Remarks. This species is known as Crambus mutabilis Clemens in the literature. Klots 
(pers. comm.) intends to put this species in a different genus. This species belongs to the 
genus Pediasia and is so treated in this paper. Bleszynski (1959) cites this species as 
Pediasia mutabilis. 


Thaumatopsis pexellus (Zeller) 
(Figs. 3D, 6a & b) 


Records. 1628 specimens from Brookings, Clay, Codington, Dewey, Harding, Jackson, 
Lawrence, Minnehaha, Spink counties. 

Flight period. Most recorded in early to mid-September, extreme dates 16 July to 1 
October. 


VOLUME 38, NUMBER 3 157 












































Fic. 8. Distribution data for: A—Pediasia trisecta, B—Pediasia dorsipunctella, C— 
Pediasia mutabilis, D—Thaumatopsis pexellus, E—Thaumatopsis fernaldellus, F—Eu- 
chromius californicalis. 


Remarks. Klots (pers. comm.) regards T. pexellus as very baffling in its local and 
individual variations. Males are needed for positive identification. The bipectinate an- 
tennae of the male separates it from all other South Dakota species. Females have filiform 
antennae and are indistinguishable from T. fernaldellus unless collected with males 
because of individual variations. 


Thaumatopsis fernaldellus Kearfott 
(Figs. 3E, 6c & d, 7h & i) 


Records. 967 specimens from Brookings, Codington, Corson, Dewey, Jackson, Lyman, 
Meade, Mellette, Pennington, Spink, Stanley counties. 

Flight period. Bivoltine with peaks in early June and early August, extreme dates 28 
May to 20 September. 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 4. a & b—Crambus praefectellus, ¢ & d—Crambus ainslieellus, e & {—Cram- 
bus alboclavellus, g & h—Crambus leachellus, i & j;—Crambus laqueatellus, k & 1— 
Chrysoteuchia topiaria. 


VOLUME 88, NUMBER 3 159 





Fic. 5. a—Crambus pascuellus floridus, b—Pediasia luteolella form caliginosella, 
e—P. |. form zeella, d—P. |. luteolella, e—Agriphila ruricolella, {—Crambus agitatel- 
lus, g & h—Agriphila vulgivagella, i—Pediasia luteolella. 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 6. a & b—Thaumatopsis pexellus, ¢ & d—Thaumatopsis fernaldellus, e & f— 
Pediasia dorsipunctella, g & h—Pediasia mutabilis. 


Remarks. The identification of all but 63 specimens recorded in this work could be 
questioned; however, all material was compared with the 63 specimens identified by Dr. 
Klots as T. fernaldellus. The character used in the key (dark stripe from base to apex of 
forewing) is often rubbed off or is hard to see on “worn’’ or light trap collected specimens. 
Work needs to be done on the genitalia within this genus. 


Thaumatopsis pectinifer Zeller 
(Fig. 1B-1) 
Records. 2 specimens from Jackson County dated 25 August. 


Remarks. The presence of the white scaled cubitus vein is the reason for these 2 
specimens being identified as T. pectinifer. 


VOLUME 88, NUMBER 3 161 





Fic. 7. a—Thamatopsis repandus, b—Argyria nivalis, e—Crambus perlellus inno- 
tatellus, d—Euchromius californicalis, e & f{—Pediasia trisecta, g—Microcrambus ele- 
gans, h & i—Thaumatopsis fernaldellus male, female, larval case. 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Thaumatopsis repandus Grote 
(Figs. 1B-2, 7a) 


Records. 7 specimens from Jackson County. 

Flight period. Most recorded as being collected in late August, extreme dates 7 August 
to 8 September. 

Remarks. Klots (pers. comm.) says the T. repandus material from South Dakota is a 
very good record. This western species represents an intrusive element in the population. 
This species is somewhat dimorphic in that the males have long or short pectinations of 
the antennae. Klots states the distribution for T. repandus as Rocky Mountain states 
westward. 


Thopeutis forbesellus (Fernald) 
(Fig. 1B-4) 


Records. 6 specimens from Brookings, Harding, Minnehaha counties dated 12 July 
and 13 August. 


Occidentalia comptulatalis (Hulst) 
(Fig. 1B-3) 


Records. 2 specimens from Jackson County dated 28 July. 


Euchromius californicalis (Packard) 
(Figs. 3F, 7d) 


Records. 602 specimens from Brookings, Buffalo, Dewey, Fall River, Harding, Jackson 
counties. 

Flight period. Peaks in mid-July, extreme dates 15 May to 2 October. 

Remarks. This is the only species of this genus we have collected from South Dakota; 
however, the females of this species and Euchromius ocelleus are difficult to separate. 
All males studied belong to E. californicalis. The genitalia are the distinguishing struc- 
ture between these two species. E. ocelleus has been recorded from North Dakota and 
can be found in South Dakota. 


Platytes vobisne Dyar 


Remarks. This species has been cited by Forbes in the literature as occurring in South 
Dakota. 


Argyria nivalis (Drury) 
(Figs. 1C-1, 7b) 


Records. 41 specimens from Brookings, Minnehaha counties. 
Flight period. Peaks in early July, extreme dates 18 June to 31 July. 


Eoreuma crawfordi Klots 


Remarks. Klots (pers. comm.) stated ““One specimen, which I have returned, is very 
unusual and to be cherished. It is almost certainly E. crawfordi which I named in 1970 
from Ames, Iowa, and Manitoba, Canada. But the abdomen is missing, so the determi- 
nation is not certain (it must, then be a new species if not crawfordi).” No additional 
material of this species has been collected to date. 


VOLUME 38, NUMBER 3 163 


ACKNOWLEDGMENTS 


We wish to thank Dr. Alexander B. Klots for aid in the determination of specimens 
and Dr. Edward U. Balsbaugh, Jr. for the loan of material from the North Dakota State 
University collection. This work is a cooperative effort of the South Dakota Agricultural 
Experiment Station, Brookings, South Dakota, and the Science and Education Adminis- 
tration, AR, USDA, as a result of coop agreement No. 12-14-3001-552. Approved for 
publication by the Director, Agricultural Experiment Station, South Dakota State Uni- 
versity, Brookings, as Journal Series No. 1965. 


LITERATURE CITED 


AINSLIE, G. G. 1922. Contributions to a knowledge of the Crambinae II. Crambus 
laqueatellus Clemens. Ann. Entomol. Soc. Amer. 15:125-136. 

1923a. Striped sod webworm, Crambus mutabilis Clemens. J. Agr. Res. 24: 

399-414. 

1923b. Silver-striped webworm, Crambus praefectellus Zincken, J. Agr. Res. 

24:415-425. 

1927. The large sod webworm, Crambus trisectus Walker. U.S.D.A. Tech. Bul. 











31. 

BLESZYNSKI, S. 1959. Studies on the Crambidae (Lepidoptera). Part XXII. On the 
systemical position of several North American species of the generic group Crambus 
Fab. s.]. Polskie Pismo Entomol. 29:447—467. 

BOHART, R. M. 1947. Sod webworms and other lawn pests of California. Hilgardia 17: 
267-307. 

Capps, H. W. 1966. Review of New World moths of genus Euchronius Guenée, with 
descriptions of two new species (Lepidoptera: Crambidae). Proc. U.S. Nat. Mus. 119: 
1-10. 

CRAWFORD, C. S. 1961. The bionomics of destructive microlepidoptera of grass fields. 
Ph.D. Thesis, Washington State University. 140 pp. 

HaRWOOD, R. F. 1964. Bionomics and control of insects affecting Washington grass 
seed fields. Wash. Agr. Exp. Sta. Tech. Bul. 44. 

Dominick, C. B. 1960. Control of the corn root webworm. J. Econ. Entomol. 53(4): 
670-672. 

1964. Notes on the ecology and biology of the corn root webworm. J. Econ. 
Entomol. 57(1):41—42. 

Ey, R. 1910. New Phycitinae and Crambinae (Lepidoptera: Pyralidae) Proc. Entomol. 
Soc. Wash. 12:204. 

FELT, E. P. 1894. On certain grass-eating insects. Cornell Univ. Agr. Exp. Sta. Bul. 64: 
47-102. 

FERNALD. 1885. North American Pyralidae. Can. Entomol. 17:55-58. 

1896. The Crambidae of North America. Ma. Agr. Coll. 93 pp. 

FORBES, W. T. M. 1920. Notes on the Crambinae (Lepidoptera). J. N.Y. Entomol. Soc. 
28(3—4):214-227. 

1923. The Lepidoptera of New York and neighboring states. Cornell Agr. Exp. 
Sta. Memoir 68. 729 pp. 

Grote, A. R. 1880. Crambidae. Can. Entomol. 12:15-80. 

KEARFOTT, W. D. 1905. Assiniboia micro-lepidoptera, collected by Mr. T. N. Willing. 
Can. Entomol. 37:119-124. 

1908. Descriptions of new species of North American Crambid moths. Proc. 
U.S. Nat. Mus. 25(1649):367-393. 

Kiots, A. B. 1940. North American Crambus. I. The silvery-striped species of Calli- 
fornia (Pyralididae). Bul. So. Calif. Acad. Sci. 39(1):53-70. 

1942. North American Crambus (Pyralididae). II. New species. Amer. Mus. 

Novit. 1191:17 pp. 

1961. Zoogeography in the systematics of North American Crambinae (Lepi- 

doptera:Pyralididae). XI International Kongress Fur Entomologie Wein 1960. Ed. I. 




















164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1963. Notes on Connecticut Sphagnum bog. J. N.Y. Entomol. Soc. 71:178-180. 
1967. Two new species of Crambus Fabricius from Western North America. 
(Lepidoptera: Pyralididae). J. N.Y. Entomol. Soc. 75(3):154-158. 

1968. The North American Microcrambus (Lepidoptera: Pyralididae). J. N.Y. 
Entomol. Soc. 76(1):9-21. 


MCDUNNOUGH, J. 1939. Checklist of the Lepidoptera. Mem. So. Calif. Acad. Sci. 2(1): 
171 pp. 

MILLER, H. D. O. 1940. Observations on sod webworms (Crambus spp. Lepidoptera) 
in Kansas. Trans. Kansas Acad. Sci. 43:267-281. 

Muma, M. H. & R. E. HILL. 1950. Thaumatopsis pectinifer (Zeller) injurious to corn 
in Nebraska. J. Kansas Entomol. Soc. 23(3):79-83. 








Journal of the Lepidopterists’ Society 
38(3), 1984, 165-170 


PAPILIO EURYMEDON LUCAS, 1852: A SYNONYM OF 
PAPILIO ANTINOUS DONOVAN, 1805 (PAPILIONIDAE) 


MURRAY S. UPTON 


Australian National Insect Collection, C.S.I.R.O. Division of Entomology, 
Canberra, A.C.T., Australia 


ABSTRACT. The holotype of Papilio antinous Donovan, 1805 has been recognised 
in the Macleay Museum, Sydney, Australia, and it is considered to be a senior synonym 
of Papilio eurymedon Lucas, 1852 of North America. The histories of these names and 
of the Donovan specimen are outlined and the nomenclatural problem discussed. 


Donovan (1805) illustrated and described Papilio antinous (Fig. 1) 
with the comment “We have observed this undescribed species only 
in the cabinet of Mr. Francillon. It was obtained by this gentleman 
from Dr. White, who resided for some time in New South Wales.” 
There was no mention of any type, and Donovan did not indicate how 
many specimens were before him. Although the provenance was not 
stated it was inferred from Donovan’s comment to be New South Wales, 
Australia. 

In 1818 Francillon’s collection was sold at auction (Chalmers-Hunt, 
1976). From an annotated copy of the sale catalogue in the Macleay 
Museum it is known that Alexander Macleay purchased a considerable 
proportion of it and he took it to Australia in 1825 with the rest of his 
vast collection. 

Godart (1819) followed Donovan, providing a more detailed descrip- 
tion of P. antinous and stating that the species came from New Hol- 
land. Boisduval (1832, 1836) also copied Donovan and referred to Go- 
dart. 

In 1844 Doubleday listed, without comment, Papilio antinous as a 
junior synonym of P. turnus Linn., 1771 (now P. glaucus Linn., 1758), 
a North American species. This synonymy was accepted by Doubleday 
(1846). 

On the death of Alexander Macleay in 1848 the Macleay collections 
were inherited by his son William Sharp Macleay, who continued to 
build and study them in collaboration with his cousin, William Ma- 
cleay. 

Early in 1852, both Lucas and Boisduval described Papilio eury- 
medon from California. Later that year Westwood (1852) added it to 
the list of diurnal Lepidoptera, and Gray (1853) listed it as being in 
the collections of the British Museum, London. 

In June 1863 William Macleay (1864) addressed the Entomological 
Society of New South Wales saying “that he wished to take the earliest 
opportunity in his power of pointing out an error in Doubleday and 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





PL AJ / / ‘ oo 
1 7 LEE VAMP OLS | 


Fic. 1. Donovan’s original figure of Papilio antinous. 


VOLUME 38, NUMBER 3 167 


Westwood’s Genera of Diurnal Lepidoptera, an error which seemed 
to have been adopted in all subsequent catalogues of Papilionidae. The 
Papilio antinous of Australia, which is figured in “‘Donovan’s Insects 
of New Holland’, from the unique specimen in the cabinet of W. S. 
Macleay, Esq., of Elizabeth Bay, is placed by Doubleday and West- 
wood as a synonym of Papilio turnus, a well known American But- 
terfly. He had not noticed this circumstance until a few days ago, but 
he had then compared the P. turnus with the P. antinous in Mr. 
Macleay’s collection, and found, as he had expected, that there was 
not even a resemblance between the species. 

“The P. antinous clearly belonged to the Podalirius group of Pa- 
piliones, and would no doubt be found (as our acquaintance with the 
Northern parts of Australia increased) to be, as originally stated, a New 
Holland insect.” 

In this statement William Macleay clearly demonstrates that the 
nominal species-group taxon was based on a single specimen—‘‘the 
unique specimen.” It is therefore clear that that specimen is the ho- 
lotype of Papilio antinous Donovan, 1805 under Article 73.(a)(ii) of 
the International Code of Zoological Nomenclature (1985). 

William Sharp Macleay died in 1865 and the collections were in- 
herited by William Macleay. 

No doubt as a result of William Macleay’s comments, Kirby (1871) 
listed P. antinous as being Australian. He also listed P. eurymedon 
from California. Later, in the first Australian catalogue to cover diurnal 
Lepidoptera, George Masters (1873), an associate of Macleay’s, also 
claimed antinous as an Australian species, as did Semper (1878). 

In 1887 William Macleay gave the Macleay collections to the Uni- 
versity of Sydney where a new building, the Macleay Museum, had 
been built to receive them. 

The second catalogue of Australian diurnal Lepidoptera to be pub- 
lished, Miskin (1891), placed P. antinous as “reputed to be Australian, 
but in support of which the evidence is not conclusive’; this appears 
to be the last occasion on which the name was used. Waterhouse (1908) 
made no mention of the name in his catalogue, nor did Bryk (1930), 
although he did refer to other species described by Donovan in 1805. 

In 1969 the C.S.I.R.O. Division of Entomology, as custodian of the 
Australian National Insect Collection, was asked to locate and hold on 
permanent loan all the type and similarly important material from the 
Macleay Museum. Although many types have been recognised and 
transferred to the Australian National Insect Collection, the search for 
further types continues. During this search Mr. Ted Edwards drew my 
attention to a specimen in a drawer of mixed papilionids. This speci- 
men (Fig. 2) bore the label ‘Papilio antinous Don. Australia” clearly 
written in George Masters’ handwriting. 


168 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


. 
peat, co 
“Bac * 





Fic. 2. The holotype of Papilio antinous Donovan, 1805. 


Masters was appointed curator of the Macleay collections by Wiiiiam 
Macleay in 1874 in order that these great collections could be reorga- 
nised and brought together. Unfortunately, during this work many 
specimens were neatly relabelled by Masters and the original labels 
discarded. This action has caused enormous problems in the recogni- 
tion of type material; indeed, some types may no longer be recognis- 


able. 


VOLUME 38, NUMBER 3 169 


There is no such problem with Papilio antinous, for it is clearly 
established (Macleay, 1864) that the unique specimen was in the Ma- 
cleay collections, and in view of its true identity there is no likelihood 
of further specimens having been added. 

It is therefore my opinion that the single specimen found in the 
Macleay Museum is the one referred to by William Macleay in 1863 
(Macleay, 1864) and is therefore the holotype of Papilio antinous Don- 
ovan, 1805. Examination of this specimen clearly shows it to be con- 
specific with the North American Papilio eurymedon Lucas, 1852, 
which name must now be considered a junior synonym under Article 
23 of the International Code of Zoological Nomenclature (1985). 

However, since the name Papilio antinous has not been cited in the 
literature since 1891, there would appear to be a clear case, under 
Article 79 of the International Code, to make application to the Inter- 
national Commission of Zoological Nomenclature for its suppression as 
the established stability of the name P. eurymedon would otherwise 
be threatened. However, the purpose of this paper is to establish the 
correct identity of Papilio antinous, and any application to the Com- 
mission should be done by those specialists directly affected by the 
change of name. 

Donovan’s figure (Fig. 1) agrees well with the specimen of P. anti- 
nous (Fig. 2), and the few discrepancies are easily explained by his 
careless approach to his work, detailed by Westwood (1872) and 
Waterhouse (1938). 


Synonymy 


Papilio antinous Donovan, 1805: plate 16; Godart, 1819:54; Boisduval, 1832:43 & 1836: 
331; Kirby, 1871:564; Masters, 1873:2; Semper, 1879:56; Miskin, 1891:83. 
Papilio turnus Doubleday (nec Linnaeus, 1771), 1844:16 & 1846:13. 
Papilio eurymedon Lucas, 1852:140; Boisduval, 1852:280; Westwood, 1852:529; Gray, 
1858:24; Kirby, 1871:565. 


ACKNOWLEDGMENTS 


I wish to acknowledge the help of Ted Edwards (C.S.I.R.O. Division of Entomology, 
Canberra) in drawing my attention to the specimen, confirming the identification and 
for commenting on the manuscript. I also wish to thank Dr. D. S. Horning of the Macleay 
Museum for his advice and John Green and Alan Edward (C.S.I.R.O. Division of Ento- 
mology, Canberra) for the photography. 


LITERATURE CITED 


BOISDUVAL, J. B. A. D. DE. 1832. Voyage de decouvertes de |’Astrolabe etc. execute 
par ordre du Roi, pendant Les annees 1826-1829, sous le commandement de M. J. 
Dumont d’Urville. Faune Entomologique de l’Ocean Pacifique. Pt. 1. J. Tastu, Paris. 
Pp. i-iv, 5-267. 

1836. Histoire Naturelle des Insectes. Species General des Lépidoptéres. Vol. 1. 

Roret, Paris, Pp. i—xii, 1-690. 





170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1852. Lépidoptéres de la Californie. Ann. Soc. Entomol. Fr. (2nd series.) Vol. 
10:280-281. 

BrYK, F. 1930. Papilionidae II (Papilio). In Lepidopterorum Catalogus, Vol. 24, Pars 
37. Pp. 57-510. 

CHALMERS-HUNT, J. M. 1976. Natural History Auctions 1700-1972. Sotheby Park Ber- 
net, London. Pp. i-xii, 1-189. 

DONOVAN, E. 1805. An epitome of the Natural History of the Insects of New Holland, 
New Zealand, New Guinea, Otaheite, and other islands in the Indian, Southern, and 
Pacific Oceans: including figures and descriptions of one hundred and fifty-three 
species of the more splendid, beautiful, and interesting insects, hitherto discovered 
in those countries, and which for the most part have not appeared in the works of 
any preceding author. Donovan, London. Pp. i-iv, 41 plates. 

DOUBLEDAY, E. 1844. List of the Specimens of Lepidopterous Insects in the Collection 
of the British Museum, Part 1. Brit. Mus. London. Pp. i-iv, 1-150. 

(1846). In Doubleday & Westwood, 1846. The Genera of Diurnal Lepidoptera: 
comprising their generic characters, a notice of their habits and transformations, and 
a catalogue of the species of each genus. Vol. 1. Longman, Brown, Green and 
Longmans, London. Pp. 7-18. 

GopakT, J. B. (1819). In Latreille & Godart, 1819. Histoire Naturelle. Entomologie ou 
Histoire Naturelle des Crustaces, des Arachnides et des Insectes, p. 54. In Encyclo- 
pedie Methodique, 9. Agasse, Paris. Pp. 1-328. 

Gray, G. R. 1853. Catalogue of Lepidopterous Insects in the Collection of the British 
Museum. Part I. Papilionidae. 1852. Brit. Mus. London. Pp. i-iii, 1-84. 

KirnBy, W. F. 1871. A Synonymic Catalogue of Diurnal Lepidoptera. J. van Voorst, 
London. Pp. i—vii, 1-690. 

Lucas, P. H. 1852. Description de nouvelles espéces de Lépidoptéres appartenant aux 
collections entomologiques de Musée de Paris. Revue et Mag. Zool. (2)4. Pp. 140- 
141. 

MACLEAY, W. 1864. Proceedings of the Entomological Society of New South Wales. 
Trans. Entomol. Soc. N.S.W. 1(2):vi-xxii. 

MasTERS, G. 1878. Catalogue of the described Diurnal Lepidoptera of Australia. Mas- 
ters, Sydney. Pp. i-iv, 1-24. 

MISKIN, W. H. 1891. A synonymical Catalogue of the Lepidoptera Rhopalocera (but- 
terflies) of Australia with full bibliographical reference; including descriptions of 
some new species. Ann. Queensl. Mus. 1:1-98. 

SEMPER, G. 1879. Beitrag zur Rhopalocerenfauna von Australien. J. Mus. Godeffroy. 
14:1-58. 

WATERHOUSE, G. A. 1903. Catalogue of the Rhopalocera of Australia. Mem. N.S.W. 
Nat. Club 1:37-88. 

1938. Notes on Jones’ Icones (Lepidoptera). (With footnotes and Appendix by 
Sir Edward B. Poulton). Proc. R. Entomol. Soc. Lond. (A), 13:9-17. 

WESTWOOD, J. O. (1852). In Doubleday & Westwood, 1852. The Genera of Diurnal 
Lepidoptera: comprising their generic characters, a notice of their habits and trans- 
formations, and a catalogue of the species of each genus. Vol. 2. Longman, Brown, 
Green and Longmans, London. Pp. 503-534. 

1872. Descriptions of some new Papilionidae. Trans. Entomol. Soc. Lond. 1872. 

(2.) Pp. 97-98, 104-110. 














Journal of the Lepidopterists’ Society 
38(3), 1984, 171-175 


HAMADRYAS IN THE UNITED STATES (NYMPHALIDAE) 


DALE W. JENKINS! 
3028 Tanglewood Drive, Sarasota, Florida 33579 


ABSTRACT. Seven species of Hamadryas have been collected in the United States. 
Species included in a recent checklist are: H. amphinome mexicana (Lucas), and H. 
feronia farinulenta (Fruhstorfer). H. februa gudula (Fruhstorfer) should be changed to 
H. februa ferentina (Godart). Additions to this list for the United States are: H. am- 
phichloe diasia (Fruhstorfer), H. guatemalena marmarice (Fruhstorfer), H. atlantis le- 
laps (Godman & Salvin), and H. iphthime joannae Jenkins. Species unsubstantiated are: 
H. fornax fornacalia (Fruhstorfer) and H. ferox (Staudinger) (correct name is H. am- 
phichloe ferox (Staudinger), and they should probably be deleted from the previous list. 
H. guatemalena (Godman and Salvin, 1883) (nec. Bates) is based on misidentifications 
of H. feronia in Texas. 


The neotropical genus Hamadryas known for many years as Age- 
ronia is a taxonomically confused genus of butterflies that has been in 
great need of revision. A critical revision of the genus has been com- 
pleted (Jenkins, 1983) so that it is now possible to accurately identify 
the Hamadryas of the United States. This revision is based on exami- 
nation of over 9000 specimens including 53 types in 30 major museums 
and collections, and on collecting and field studies by the author in 20 
countries. Of the 100 taxa named, only 20 species and 21 subspecies 
are recognized. Keys to male and female adults, male genitalia, and 
descriptions and distributions are included in the above mentioned 
revision. 

Hamadryas spp. have been difficult to determine because needed 
identification characters and keys have not been published previously, 
there is much confusion due to a plethora of synonyms created by 
Fruhstorfer, Bryk and others, and misidentifications of figures exist in 
many books. Numerous figures in Fruhstorfer in Seitz (1916) are er- 
roneous. Klotz (1951, pl. 18) and Howe (1975, pl. 15) both identify 
figures of H. februa ferentina as H. feronia. In the recent book ‘“‘Au- 
dubon Society Field Guide to North American Butterflies’ by Pyle 
(1981) all Hamadryas pictures are misidentified. Picture 758, “white 
skirted calico,” is identified as Hamadryas feronia, but it is actually 
H. februa ferentina, the most frequently collected species of Hama- 
dryas in the United States. The common name designated for H. fer- 
onia is unfortunate since H. feronia farinulenta from Texas and 
Mexico has dark buff to light ochre ventral hind wings. Picture 759, 
“yellow-skirted calico,’ misidentified as Hamadryas fornax is actually 
H. guatemalena marmarice (taken in Mexico). I have found no valid 


1 Research Associate, Allyn Museum of Entomology, Sarasota, Florida. 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


or confirmed records of H. fornax in the United States. The common 
name “‘yellow-skirted calico’ is not descriptive since the ventral hind 
wings of H. fornax are dark mustard to orange in color. Hamadryas 
are often known as “crackers” because of the crackling or clicking 
noise they make in flight. Some species are called “calicoes’” because 
of the complex mosaic pattern on the wings that provides camouflage 
when they alight with wings outspread on tree trunks. 

All available specimens of Hamadryas collected or purported to 
have been collected in the United States have been critically examined 
and determined. 

A list of valid collection records for the United States is presented 
which includes seven species, four of which are new for the United 
States and not reported in Miller and Brown (1981), plus nomencla- 
torial changes. Three species previously reported by error for the United 
States have been based on misidentifications. 

The following nomenclature follows Jenkins (1983). 


A. Valid Records for the United States 


1. Hamadryas amphinome mexicana (Lucas, 1853) 

Specimens identified: (photograph) Texas, Hidalgo Co., Bentsen-Rio Grande Val- 
ley State Park. 1 6 (fresh), 3 Sept. 1972, Leg. W. W. McGuire (McGuire & Rickard, 
1974). This specimen in the McGuire collection was photographed and published by 
Kendall (1974) showing dorsal and ventral views permitting identification of the 
subspecies as mexicana. It is the only positive collection record known to me in the 
United States and is almost certainly a chance migrant or stray. The nearest known 
record is at Tamazunchale, Mexico, over 500 km south. However, this subspecies 
became established in western Cuba and was commonly collected in 1934 and 1935. 

In Miller and Brown (1981), Catalogue No. 685a. 

2. Hamadryas februa ferentina (Godart, [1824]) 

Specimens identified: “Texas,” 1 2 labelled “type” of Ageronia februa gudula 
(Fruhstorfer, 1916) in the British Museum (Natural History). I have examined and 
photographed this type, and it is a dark female of H. februa ferentina. The original 
description states that it comes from western Mexico. Texas, Hidalgo Co., Bentsen- 
Rio Grande Valley State Park, 2 92 24 Aug. 1969, Leg. M. A. Rickard, in Amer. Mus. 
Nat. Hist. Coll.; Texas, Pharr, 1 2 Leg. H. A. Freeman, in O. Buckholz Coll., in 
Amer. Mus. Nat. Hist. Coll.; Texas, Hidalgo Co., Bentsen-Rio Grande Valley State 
Park, 1 2 (worn), 1 6 24 Aug. 1969, Leg. Roy Kendall; 1 6 (fresh) 30 Aug. 1973, Leg. 
M. A. Rickard in Roy Kendall Coll.; Texas, Hidalgo Co., Loop 37, 6 mi. W of Mission 
(fresh), 19 Oct. 1973, Leg. W. William and N. McGuire, in Roy Kendall Coll.; 
“Texas” 1 6, 1 2 in Carnegie Museum Coll.; “Texas” 1 6 Coll. T. L. Mead in Holland 
Coll., No. 299, labelled “Ageronia feronia (Linn.)” in the Carnegie Museum Coll. 

Other records: Reported from Texas by McGuire and Rickard (1974), Howe (1975), 
and others as H. februa gudula (Fruhstorfer) which is one of many synonyms of H. 
februa ferentina. This is the most commonly collected Hamadryas in the United 
States and is probably a resident or becomes established regularly in southern Texas. 
The larvae probably feed on the euphorbiaceous plant Tragia which occurs in south- 
ern Texas. 

In Miller and Brown (1981) as Hamadryas februa gudula (Fruhstorfer) No. 683a. 

3. Hamadryas feronia farinulenta (Fruhstorfer, 1916) 
Specimens identified: Texas, Hidalgo Co., Loop 37, 6 mi. W of Mission, 1 6 (fresh) 


VOLUME 38, NUMBER 3 173 


15 Jul. 1975. In Roy Kendall Coll. “Texas” labelled “A. formax” [Sic.] 1 6 in Los 
Angeles Co. Mus. Nat. Hist. 

Other records of “‘feronia’’: “This remarkable insect is said to be occasionally found 
in Texas’ Holland (1898). Texas, Pharr. “Strays” in Oct. and Nov., Klots (1951). 
Texas, Brownsville, “Strays,” Howe (1975). “Southern Texas.”’ Many reports and 
quotes with no specific data starting with Scudder (1875) need verification. The two 
male records from Texas appear to be the only valid records of this species for the 
United States. No other United States specimens have been found in any of the 
museum collections studied. (This is frequently confused with H. februa ferentina. 
See misidentified record in Holland Coll. above.) 

In Miller and Brown (1981), Catalogue No. 682a. 

4. Hamadryas amphichloe diasia (Fruhstorfer, 1916) 

Specimens identified: (Photograph). Florida, Monroe Co., Plantation Key, 5 Jul. 
1978, Leg. Paul Tuskes. Florida, Monroe Co., Key Largo, Tavernier, 16 Jul. 1978 
(sight record) Paul Tuskes. These specimens were reported as Hamadyras februa 
diasia in the 1978 Field Summary (1979). It is unknown whether they are migrants, 
temporary, or permanent residents. Tragia saxicola Small occurs in the Florida Keys 
as well as two other species of Tragia which could be host plants. H. amphichloe 
diasia occurs in Cuba, Jamaica, and Puerto Rico where it is relatively uncommon or 
rare and in Hispaniola where it is fairly common. H. amphichloe diasia has been 
called H. ferox diasia. As stated in Jenkins (1983) Ageronia amphichloe Boisduval, 
1870 was incertae sedis for over 100 years due to the poor description. I examined 
the original Boisduval type in the British Museum and found that Ageronia ferox 
tegyra Fruhstorfer, 1916 is a synonym, and H. ferox diasia becomes H. amphichloe 
diasia. (For H. “ferox’”’ records in Texas see list of unsubstantiated records below.) 

Not listed in Miller and Brown (1981). 

5. Hamadryas guatemalena marmarice (Fruhstorfer, 1916) 

Specimens identified: Texas, Hidalgo Co., Bentsen-Rio Grande Valley State Park, 
1 4 (fresh), 17 Aug. 1974, Leg. Frank Hedges, on loan to Roy Kendall Coll. (a specimen 
of this subspecies was photographed by Harry Darrow in Mexico, and published by 
Pyle [1981] as H. fornax). The Texas specimen appears to be the first and only valid 
United States record. This subspecies occurs from Tamaulipas in northeastern Mexico 
and Sonora, Mexico, to the Isthmus of Tehuantepec and Chiapas. Further details are 
being published by Kendall. 

Other records of “H. guatemalena”: Godman and Salvin (1883) confused and 
misidentified H. feronia as H. guatemalena and assumed that H. feronia reported 
for Texas by Strecker (1878) was H. guatemalena. 

Not listed in Miller and Brown (1981). 

6. Hamadryas iphthime joannae Jenkins, 1983 

Specimens identified: Texas, Burnet Co. 1 6 (fresh) Aug., William C. Wood Coll., 
in the Amer. Mus. Nat. Hist. Coll. This is northwest of Austin, Texas, and is surely 
a stray migrant since the nearest records are at Tamazunchale and Tuxpan, Veracruz, 
Mexico, about 1000 km south. This is a new record for the United States. The 
specimen had been identified previously as H. iphthime (Bates) by F. M. Brown but 
was never published. Ageronia iphthime Bates was described from a syntype from 
Bogota, Colombia, and partly from a syntype from Guatemala. I have designated 
the Bogota type as lectotype of the Panama and South American population and the 
Mexican and Central American population was described as H. iphthime joannae. 
See below for an erroneously labelled and misidentified specimen of H. iphthime 
iphthime. 

Not listed in Miller and Brown (1981). 

7. Hamadryas atlantis lelaps (Godman and Salvin, 1883) 

Specimens identified: (Photograph). Arizona, Cochise Co., Douglas, San Bernadino 
Ranch, 1 4, 14 Aug. 1976, Leg. Peter Jump. The photograph was sent by Richard A. 
Bailowitz who also reports an additional sight record. 

H. atlantis lelaps was recognized as a new subspecies by Jenkins (1983) after 


174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


collecting specimens in Sinaloa, Mexico, and comparing with the 2 type of Ageronia 
lelaps in the British Museum. Godman and Salvin in 1901 had synonymized Age- 
ronia lelaps as the 2 of Ageronia atlantis. Fruhstorfer in Seitz (1916) misidentified 
lelaps and applied the name to an undescribed Hamadryas glauconome grisea Jen- 
kins (1983) which occurs in Sonora, Mexico, and which should be collected in Arizona 
and/or New Mexico. 

This new record for the United States is a significant northward extension of range 
of H. atlantis lelaps which had previously been reported as far north as Alamos, 
Sonora, Mexico. 


B. Unsubstantiated Records 


1. Hamadryas fornax (Hubner, [1828]) 

H. fornax fornacalia (Fruhstorfer, 1907) is an uncommon and locally occurring 
subspecies that is definitely known as far north as near Tampico, Mexico, about 400 
km south of Texas. H. fornax was reported from southern Texas by Scudder (1875) 
and quoted by many authors. Holland (1898) states “A. fornax is reported only from 
the hotter parts of Texas.’’ I examined the Holland Collection in the Carnegie Mu- 
seum which revealed a specimen labelled Ageronia fornax Hubn. in purple ink, 
probably in Holland’s handwriting. It is labelled “Texas or N.M./Coll. T. L. Mead/ 
Holland Coll.” It is a misidentified male of Hamadryas iphthime iphthime which is 
found from Costa Rica to southern South America. Klots (1951) reported it from 
Pharr, Texas, in Oct., and Howe (1975) in the Brownsville area. Pyle (1981) misi- 
dentified and published a picture of H. guatemalena marmarice from Mexico as H. 
fornax for Texas. Despite all these publications I have not been able to confirm any 
valid specimens in over 30 major museums and private collections examined. 

In Miller and Brown (1981), listed as Hamadryas fornax fornacalia, No. 681a. 

2. Hamadryas amphichloe ferox (Staudinger, [1886]) 

H. amphichloe ferox is a rare species that has been reported from Texas. It occurs 
in Venezuela and Colombia and has been reported from Central America. I have 
studied and photographed the single holotype male of H. ferox fictitia (Fruhstorfer, 
1916) from “Mexico” in the British Museum (Natural History). This is a synonym 
of H. amphichloe ferox. The locality record of Mexico is probably fictitious. I have 
examined two other old specimens of H. amphichloe ferox from Central America 
which are probably mislabelled: 1 6 from “Guatemala,” U.S. National Museum Coll., 
and 1 6 from “Panama” in the Strecker Coll. at the Allyn Museum, neither with 
specific localities. Klots (1951) states “There is also a vague record of H. ferox 
Staudinger from southern Texas which I have been unable to verify.” Hoffman 
(1940) also was dubious of H. ferox fictitia and knew of no locality in Mexico where 
it occurred. 

In Miller and Brown (1981), listed as Hamadryas ferox (Staudinger), No. 684. An 
asterisk was used to indicate that it was of doubtful occurrence in the United States. 
This record should be deleted from the list. 

8. Hamadryas guatemalena (Bates, 1864) 

Godman and Salvin (1883) include the record of H. feronia from southwest Texas 
by Scudder (1875) and other authors as H. guatemalena. This was due to their 
confusion of these two species. They considered H. feronia as occurring from Pan- 
ama south and that H. guatemalena occurred in Central America and Mexico. This 
was based on misidentifications, and no known specimens were then available from 
Texas to confirm H. guatemalena. 


ACKNOWLEDGMENTS 


I would like to thank Roy O. Kendall for sending several Texas specimens of Hama- 
dryas for determination and for comments on the manuscript, and to curators of nu- 
merous museums and private collections for permission to review their Hamadryas. I 
greatly appreciate the kind help and valuable comments of Dr. Lee D. Miller, Jacqueline 


VOLUME 38, NUMBER 3 175 


Y. Miller, and Dr. Arthur Allyn and for continuing use of the collections and facilities 
of the Allyn Museum of Entomology, Fla. State Museum. 


LITERATURE CITED 


GODMAN, F. D. & O. SALVIN. 1883. Biologia Centrali-Americana. Insecta. Lepidoptera- 
Rhopalocera. London 1:273-274. 

HOFFMANN, C. C. 1940. Catalogo sistematico y zoogeografico de los lepidopteros mex- 
icanos. Primiera parte. Papilionoidea. An. Inst. Biol. Mexico 11:639-739. 

HOLLAND, W. J. 1898. The Butterfly Book. Doubleday and McClure, New York. 
382 pp. 

HoweE, W. H. (Ed.). 1975. The Butterflies of North America. Doubleday and Co., Inc. 
Garden City, N.Y. 633 pp. 

JENKINS, D. W. 1983. Neotropical Nymphalidae. I. Revision of Hamadryas. Bull. Allyn 
Mus. 81:1-146. 

KENDALL, R. O. 1974. Confirmation of Rhopalocera-Pieridae, (Nymphalidae) previ- 
ously recorded for Texas and the United States. J. Lepid. Soc. 28(3):249-252. 

Kiots, A. B. 1951. A Field Guide to the Butterflies of North America, East of the 
Great Plains. Houghton Mifflin Co., Boston. 349 pp. 

McGuIrE, W. W. & M. A. RICKARD. 1974. An annotated checklist of butterflies of 
Bentsen-Rio Grande Valley State Park and vicinity. Texas Parks and Wildlife Dept. 
Mimeo. 23 pp. 

MILLER, L. D. & F. M. BRowNn. 1981. A Catalogue/Checklist of the Butterflies of 
America North of Mexico. Lepid. Soc. Mem. No. 2, 280 pp. 

PyLE, R. M. 1981. The Audubon Society Field Guide to North American Butterflies. 
Alfred A. Knopf, New York. 916 pp. 

SCUDDER, S. H. 1875. Historical sketch of the generic names proposed for butterflies. 
Proc. Amer. Acad. Arts and Sci. 10:109. 

SEITZ, A. 1916. The Macrolepidoptera of the World. 5. Ageronia. 537-545. 

STRECKER, F. H. H. 1878. Butterflies and Moths of North America. 283 pp. 

WINTER, W. D. (Ed.). 1979. Field Summary for 1978. Zone 6. South Florida, News 
Lepid. Soc. 2:11. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 176-178 


A SEX PHEROMONE IN THE CALIFORNIA OAKWORM 
PHRYGANIDIA CALIFORNICA PACKARD (DIOPTIDAE) 


MICHAEL E. HOCHBERG AND W. JAN A. VOLNEY 


Division of Entomology and Parasitology, 
University of California, Berkeley, California 94720 


ABSTRACT. California oakworm (Phryganidia californica) virgin females confined 
to sticky traps attracted significantly more males than unbaited control traps. This dem- 
onstrates the presence of a sex attractant in this species. 


The California oakworm (COW), Phryganidia californica Packard, 
the only species of the family Dioptidae in America north of Mexico, 
is a major defoliator of oaks in California (Essig, 1958; Brown & Eads, 
1965). Previous studies (Harville, 1955; Sibray, 1947) have shown that 
COW populations erupt sporadically, but the causes of these eruptions 
are presently unknown. Attractive pheromones, should they exist, could 
provide a means of detecting sparse populations and incipient out- 
breaks and determining the distribution of this species (Daterman, 
1978; Cardé, 1979). 

Here we report results that indicate the presence of a female pro- 
duced sex pheromone in this species. 


MATERIALS AND METHODS 


The study was carried out in October 1982 in a ca. %4 hectare stand 
of California live oaks (Quercus agrifolia Neé) on the University of 
California campus, Berkeley. Adults used in these trials were field 
collected pupae which were confined individually to 90 x 23 mm shell 
vials plugged with cotton wool. The insects were reared under a natural 
photoperiod in the laboratory and allowed to emerge in these vials. 

Pherocon 1C® (Zoecon Corp., Palo Alto, CA) sticky traps were used 
in all trials. Traps were baited by confining one virgin female to a 
cylindrical (6 x 12 mm) steel mesh cage suspended from the trap roof. 
A 5 ml water vial plugged with cotton was also included in cages. 
Control traps were each fitted with a cage containing a water vial but 
no female. Traps were placed in arbitrarily selected California live 
oaks, hung between 2 m and 4 m above ground, and at least 8 m apart. 

In the first trial, 10 traps were baited with females which had emerged 
from pupae 0-24 hours prior to the experiment. Ten control traps were 
also deployed. In the second trial, eight traps were baited with females 
which had eclosed 12-24 hours prior to the experiment, 12 with fe- 
males that eclosed 24-36 hours prior to trap placement and 10 unbaited 
traps served as controls. In each trial traps were examined 24 hours 
after they were deployed. 


VOLUME 38, NUMBER 3 477 


TABLE 1. Male moths caught by traps baited with virgin females and unbaited con- 
trols. 


Female age Number Standard 
(hours)* of traps Mean catch deviation Range 
Trial I Control a 10 0.1 0.3 0-1 
0-24 b 10 23.7 29.3 0-70 
Trial II Control a 10 0.1 0.3 0-1 
12-24 b 8 66.4 41.4 0-114 
24-36 b 12, 55.4 46.5 0-126 


; + cae trials, treatments (female age at trap deployment) followed by the same letter are not significantly different 
a= 0.05). 


RESULTS 


The number of males caught in each trap for a particular trial was 
ranked and these data were analyzed by means of the Mann-Whitney 
test (Conover, 1971, p. 224). In each trial, traps baited with virgin 
females caught a significantly greater number of males than unbaited 
controls (Table 1). In the second trial, although the number of male 
moths caught in traps baited with the younger females caught more 
males than the older females, this difference was not significant. In 
both trials, some females failed to attract moths (Table 1). 

Since baited traps caught a significant number of moths and all 
COW moths trapped were males, these results demonstrate the pres- 
ence of a sex attractant in this species. Thus, attempts to extract, isolate, 
and identify the active secretions seem justified, not simply because of 
the economic importance of this species, but also because of its unique 
position among North American Lepidoptera. 


ACKNOWLEDGMENTS 


We thank A. M. Liebhold, J. E. Milstead and D. L. Wood for reviewing this manu- 
script. Work leading to this manuscript was conducted under Hatch project 3689-H in 
the California Agricultural Experiment Station entitled Biology and Dynamics of Forest 
Insect Populations. 


LITERATURE CITED 


BROWN, L. R. & C. O. Eaps. 1965. A technical study of insects affecting the oak tree 
in southern California. Calif. Agr. Exp. Sta. Bull. 810. 105 pp. 

CaRDE£, R. T. 1979. Behavioral responses of moths to female-produced pheromones 
and the utilization of attractant baited traps for population monitoring. In R. L. 
Rabb & G. G. Kennedy (eds.). Movement of Highly Mobile Insects: Concepts and 
Methodology in Research. N.C. State Univ., Raleigh, N.C. 456 pp. 

CONOVER, W. J. 1971. Practical Nonparametric Statistics. John Wiley and Sons Inc., 
New York. 462 pp. 

DATERMAN, G. E. 1978. Monitoring and early detection. In M. H. Brookes, R. W. Stark 
& R. W. Campbell (eds.). The Douglas-Fir Tussock Moth: A Synthesis. USDA, For. 
Serv. Serv. Tech. Bull. 1585:99-202. 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Essic, E. O. 1958. Insects of Western North America. The Macmillan Co., New York. 


1050 pp. 
HARVILLE, J. P. 1955. Ecology and population dynamics of the California oak moth, 


Phryganidia californica Packard (Lepidoptera: Dioptidae). Microentomol. 20:83- 


166. 
SIBRAY, W. S. 1947. Bionomics of the California Oak Moth. Masters Thesis, University 


of California, Berkeley. 80 pp. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 179-185 


POPULATION BIOLOGY OF THE GREAT PURPLE 
HAIRSTREAK, ATLIDES HALESUS, 
IN TEXAS (LYCAENIDAE) 


PAUL L. WHITTAKER 


Department of Zoology, University of Texas, 
Austin, Texas 78712 


ABSTRACT. The population biology of the great purple hairstreak, Atlides halesus 
Cramer, was studied at two sites in Texas. A. halesus feeds on mistletoe (Phoradendron 
tomentosum Engleman) and prefers the younger, non-woody parts of the plant. It is 
most common in late spring and prefers isolated host plants for oviposition at the southern 
site. Hymenopteran parasitoids are the major source of mortality, emerging from eggs, 
third instar larvae and pupae. 


Atlides halesus Cramer is the largest widely distributed lycaenid 
butterfly in the United States. It ranges northward from Mexico to 
New York and Oregon, and is locally common in most of the southern 
states (Howe, 1975). Larvae feed on different species of mistletoe 
(Phoradendron, Loranthaceae). Haskin (1933) has described the life 
history of A. halesus in Florida. This article describes its population 
biology at two locales in Texas and includes information on time of 
development, phenology, response to host density, use of different plant 
parts and sources of mortality. American mistletoe (Phoradendron to- 
mentosum Engl.) is the host of A. halesus in most parts of Texas. P. 
tomentosum is a widely distributed hemiparasitic shrub which infects 
a variety of deciduous trees. In central Texas, it is most common on 
ulmaceous hosts (Ulmus crassifolia, Celtis spp.), while further south it 
has apparently adapted to mesquite (Prosopis glandulosa Torrey, Le- 
guminoseae). 


METHODS 


Two field sites were used, one for rearing caterpillars and the other 
for field observations. The Brackenridge Field Station of the University 
of Texas is located on the Colorado River, about two miles north of 
Town Lake in Austin, Texas. Elm (Ulmus crassifolia) and hackberry 
(Celtis spp.) are the major hosts of P. tomentosum. A. halesus is com- 
mon in late spring, with adults sometimes found in groups of three or 
four on flowers. Adults are occasionally found flying in mid-winter in 
the Austin area. Three A. halesus larvae were collected at the Brack- 
enridge Station and brought back to the lab for rearing, on 12 April 
1979 (#1) and 16 April 1979 (#2, #3). The larvae were fed fresh P. 
tomentosum leaves and weighed daily on a CAHN Electrobalance 
until pupation. 

The Chaparral Wildlife Management Area (CWMA) of the Texas 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Size classes of leaves and stems. 


Stems Leaves 
Size class (diameter) (mm) (surface area/leaf) (mm?) 
] Greater than 10 Greater than 500 
2 5-10 300-500 
3 2.5-5.0 100-300 
4 0-2.5 0-100 


Parks & Wildlife Department was used for field observations. The 
CWMA is about 60 miles north of the Mexican border at Laredo, with 
desert grassland vegetation dominated by mesquite (Whittaker et al., 
1979). Mesquite is the only major host of P. tomentosum. Most of the 
mistletoe is on older trees found in disturbed areas. A total of 375 P. 
tomentosum at four sites were individually marked in 1980 and cen- 
sused once a month between March 1981 and October 1981 (insect 
activity was low during the winter months so no data were collected). 
Each of the four sites had a windmill or water pump and a large tank 
for holding water. They were surrounded by fencing, and brush inside 
the fenced area had been periodically removed, leaving a few large 
trees, most of which were heavily infested with P. tomentosum. For 
each marked P. tomentosum, I recorded the diameter of the host 
(mesquite) tree, the distance of the tree from the nearest holding tank 
and the number of P. tomentosum on the tree. P. tomentosum plants 
were chosen to include a variety of host diameters, distances from 
water and densities of infestation (see Whittaker, 1982 for details). 
Plants were searched for A. halesus as thoroughly as possible. Because 
A. halesus larvae are very cryptic, some caterpillars were undoubtedly 
missed, especially on large, leafy mistletoe plants. For every A. halesus 
encountered, I recorded the stage of development (egg or larval instar 
number), the number of the host plant, and the part of the plant it 
was on (leaf, stem, or inflorescence). Leaves and stems were further 
divided into four size categories (Table 1). Data were analyzed to 
determine relative preference by A. halesus for P. tomentosum at 
different distances, densities and host diameters, and for different parts 
of the plant. More information on data collection and analysis is con- 
tained in Whittaker (1982). 


RESULTS 


Eggs are scattered over the host plant by the ovipositing female. As 
many as 22 were found on one plant, but they are not laid in clusters. 
Sometimes eggs are found on the branch of the mistletoe’s host tree 
near the site of infection. Larval eclosion from the egg leaves an open- 


VOLUME 38, NUMBER 3 181 


900 


400 


300 


200 


Weight (mg) 


12 14 I6 18 20 22 24 
April 


Fic. 1. Growth curves for three A. halesus larvae. 


ing on the top of the egg and may be distinguished from parasitoid 
emergence, which leaves an opening on the side of the egg. The egg 
parasite is presumably a chalcidoid or proctotrupoid wasp but was not 
identified. The eggs have a hard outer surface and are easily noticeable. 
According to Haskins (1933), the duration of the egg stage is about 
seven days. 

Larvae are green and darken with age. They are similar to a mis- 
tletoe leaf in color and texture, and extremely cryptic. Of the larvae 
collected at the Brackenridge site, one grew from 124 mg to 392 mg 
in four days before pupating, one grew from 11.7 mg to 281 mg in 
eight days and one increased from 15.7 mg to 553 mg in nine days 
(Fig. 1). Haskin’s (1933) larvae spent about 20 days between hatching 
and pupation. Pupae are dark brown and are sometimes found at the 
base of trees on which mistletoe grows (Emmel & Emmel, 1973). The 
minimum duration of the pupal stage seems to be about 16 days (Has- 
kin, 1933) (no records were kept of pupal eclosion by the three cater- 
pillars from Brackenridge). Fig. 2 shows the phenology of A. halesus 
larvae at CWMA in 1981. Caterpillars are most common in late April 
but remained common through early June and were found during all 
censuses. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ox2 


0.| 


Avg. Number of Insects /Plant 


PUN =I TEN eV TENS S/S SIN Sane 
Date 
Fic. 2. Phenology of A. halesus at CWMA. 


Fig. 3 shows the percent of observations of A. halesus larvae on 
different parts of the plant. Young (type 4) leaves were fed on most 
often, followed by fairly young (type 3) leaves, young (type 4) stems 
and fairly old (type 2) leaves. Early instar caterpillars fed on young 
leaves or rasped the surface of older leaves; late instars can chew through 
the older leaves. A. halesus larvae occasionally defoliated entire plants, 
leaving only woody (type 1 and type 2) stems remaining. Fig. 4 shows 
the observed number of larvae found on mistletoe plants growing at 
different densities divided by the expected number, based on a uniform 
distribution (Whittaker, 1982, p. 73). There were more A. halesus 
caterpillars than expected at low density infestations. The distribution 
of eggs showed a similar pattern; so, this distribution reflects preference 
by the ovipositing female and not differential mortality. A. halesus was 
also relatively more common on small diameter mesquite trees and far 
away from water tanks (Whittaker, 1982), both of which were corre- 
lated with low mistletoe density. 


yis0 
Oo €& 
3-2 40 
3 5 20 
x 
O 
Meanie ene 2 NTN te Oh» UR Crain 
Stems Leaves Inflor. 


Fic. 3. Feeding preferences for different parts of the host plant. 


VOLUME 38, NUMBER 3 183 


3.00 
2.00 


|.00 


354-6 -9- 15° 16-30 3l* 


Number of Mistletoe / Mesquite Tree 
Fic. 4. Response of A. halesus to host plant density. 


Observed Number of Insects 
Expected Number 


Fig. 5 shows the age structure of A. halesus found on censused 
plants. Eggs are probably over-represented, because they aren’t cryptic 
like the larvae. Parasitism by an unidentified hymenopteran is a major 
source of egg mortality. First and second instar larvae weren’t readily 
distinguishable, so they are grouped together. A braconid parasitoid 
(Apanteles sp.) emerged from many third instar larvae, leaving behind 
a cocoon underneath the dorsal integument of the caterpillar, and a 
large chalcidoid (Metadontia amoena Say) emerged from some pupae. 
Starvation following host plant defoliation was another source of mor- 
tality. Fig. 5 reflects the high incidence of mortality due to parasitism 
in the egg stage, third larval instar and pupae. Adults may be under- 
represented, because unlike the larvae, they don’t spend all their time 
on the host plant and were only occasionally observed during ovipo- 
sition. 


DISCUSSION 


Atsatt (1981) has hypothesized that selection for ““enemy-free space” 
has been responsible for many of the radiations in host plant use within 
the Lycaenidae. Lycaenids have independently switched hosts to mis- 
tletoes (Loranthaceae, Viscaceae) several times. Many mistletoes are 
frequented by ants, which sometimes tend homopteran populations for 
nectar and protect lycaenid caterpillars from parasites and predators. 
P. tomentosum in Texas supports several species of host-restricted 
aphids and scale insects, all of which are tended by ants (Whittaker, 
1982). No interactions between A. halesus larvae and any of the ant 
species were observed in the course of my study, and I suggest that 
the shift of Atlides onto mistletoe was a result of the nutritive qualities 
of the plant and not protection by ants. The remarkable cryptic col- 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 









Adults: 3 

5" instar: 15 
4" instar: 14 
3" instar:57 


Ist & 24 instars: 68 






Eggs:489 





Fic. 5. Numbers of A. halesus of different ages. 


oration of A. halesus larvae may offer protection against avian pred- 
ators, but the high rates of parasitism indicate that it is not effective 
against hymenopterans. The noctuid moth Emarginea (Cyathissa) per- 
cara Morrison is preyed on heavily by hemipterans (Largus cinctus 
Herrich-Schaeffer, Largidae and Podisus acutissimus Stal, Pentatom- 
idae) and attacked by at least one of the ant species which frequent 
P. tomentosum. E. percara feeds on the same parts of the mistletoe 
plant as A. halesus but is much more abundant in the Chaparral area 
and is more likely to be found on older, high density mistletoe infes- 
tations (Whittaker, 1982). E. percara defoliation is a major source of 
mortality for P. tomentosum. A. halesus also kills mistletoe plants and 
also threatens the resource base for the ants’ homopteran herds, but its 
thick integument probably protects it from attack by both ants and 
hemipterans (Malicky, 1970). Many butterfly species have now been 
found to oviposit preferentially on isolated host plants (Courtney & 
Courtney, 1982). In the case of A. halesus, I believe it is a mechanism 
for reducing competition with the voracious and destructive E. per- 
cara, which is not a strong flier as an adult and probably has trouble 
reaching isolated mistletoe plants. 


ACKNOWLEDGMENTS 


I thank Don Harvey and John Rawlins for their comments on this manuscript, Phil 
DeVries for help with field work, and Bill Brummell and Ernie Davis for letting me use 
the Chaparral Wildlife Management Area as a study site. 


LITERATURE CITED 


ATSATT, P. R. 1981. Lycaenid butterflies and ants: Selection for enemy-free space. 
Amer. Nat. 118:638-654. 

COURTNEY, S. P. & S. COURTNEY. 1982. The “edge effect” in butterfly oviposition: 
Causality in Anthocaris cardamines and related species. Ecol. Entomol. 7:181-187. 

EMMEL, T. C. & J. F. EMMEL. 1973. The Butterflies of Southern California. Natural 


VOLUME 38, NUMBER 3 185 


History Museum of Los Angeles County, Science Series #26. Los Angeles, Calif. 148 


pp. 
HASKIN, J. R. 1933. Thecla halesus, its life cycle and habits. Entomol. News 44:72-74. 
Howe, W. H. 1975. The Butterflies of North America. Doubleday, New York. 633 pp. 
Ma.icky, H. 1970. New aspects on the association between lycaenid larvae and ants. 
J. Lepid. Soc. 24:190-202. 
WHITTAKER, P. L. 1982. Community ecology of Phoradendron tomentosum in south- 
ern Texas. Ph.D. Dissertation, Univ. of Texas, Austin. 122 pp. 
WHITTAKER, R. H., L. E. GILBERT & J. H. CONNELL. 1979. Analysis of a two-phase 
pattern in a mesquite grassland, Texas. J. Ecol. 67:935-952. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 186-191 


FORAGING BEHAVIOR OF TAWNY EMPEROR CATERPILLARS 
(NYMPHALIDAE: ASTEROCAMPA CLYTON) 


NANCY E. STAMP! 


Department of Zoology, University of California, 
Davis, California 95616 


ABSTRACT. Tawny emperor caterpillars moved up to 3 m to new feeding sites, 
passing by numerous leaves in the process. These cryptic larvae molted on the underside 
of leaves or between leaves they tied together before feeding at the new sites. 


Cryptic and thus presumably palatable caterpillars may avoid their 
natural enemies by feeding on the underside of leaves, foraging at 
night, commuting to and from feeding sites, moving some distance 
between feeding bouts and cutting off leaf remains after feeding on 
leaves (Heinrich, 1979). This appears to be a consequence of birds 
learning to forage preferentially on plants with caterpillar-damaged 
leaves (Greenberg & Gradwohl, 1980; Heinrich & Collins, 1983). Para- 
sitoids also use damaged leaves and frass to locate caterpillars (Sato, 
1979). 

Tawny emperor caterpillars (Asterocampa clyton flora (Edwards): 
Nymphalidae) are of particular interest here, because they aggregate 
in the early instars, in contrast to most cryptically-colored caterpillars. 
By aggregating, these caterpillars may cause considerable leaf damage 
at a feeding site and thus, draw attention to themselves in a way that 
early instars of solitary, cryptic larvae would not. The objective of this 
study was to examine the foraging behavior of early instar tawny em- 
peror caterpillars. 


METHODS 


The caterpillars were observed on hackberry trees (Celtis laevigata 
Willdenow) at the University of Florida (Gainesville) in fall 1981. Egg 
clusters were located on the underside of leaves on the distal portion 
of major branches. Larval aggregations were followed by placing la- 
beled bands on leaf petioles of occupied leaves. Each day caterpillars 
were censused by searching leaves of the main branch with the egg 
cluster. During the molting periods these censuses provided a reliable 
record of total surviving larvae. However, during feeding bouts cat- 
erpillars were often moving back and forth along the branches and 
thus difficult to census accurately. Consequently, only larvae on leaves 
were censused. Leaf tissue eaten by the caterpillars was estimated to 


Present address: Department of Biological Sciences, SUNY, Binghamton, NY 13901. 


VOLUME 38, NUMBER 3 187 


Total/ 

day 130 127 110 90 8 94 77 5| 87 73 7I 7I 7 7 55 64 
Branch 

tip 94 77 5| —5 


Group A . . 2 *~3 


Position along branch in cm 








1 
-' leaves | : ' leaves | Se 
I ' passed : : assed . 
3 33 2 Es oe Shae =e 
2 XX s 


3 


Fic. 1. Foraging behavior of tawny emperor caterpillars, from 24 September through 
9 October 1981. EC indicates the position of the egg cluster on the branch. Larval 
numbers per leaf are shown. Arrows indicate periods of larval movement. 


Cumulative leaves 
eaten between molts 


the nearest tenth and then averaged for number of leaves eaten per 
aggregation per observation. 


RESULTS AND DISCUSSION 


After hatching, the caterpillars fed on that same leaf and occasion- 
ally on adjacent leaves. For example, group A ate 85% of the leaf tissue 
of two of three adjacent leaves. Group B fed on four adjacent leaves, 
removing 60% of the leaf material. Five to six days later the larvae 
moved up to 1.2 m to a new site, passing by numerous leaves and 
presumably suitable food along the way (Figs. 1 & 2). Reaggregation 
at the next site took one to two days. Then larvae molted on the 
underside of a leaf or between adjacent leaves, often tying the edges 
of the leaf or leaves around them. After molting the larvae fed on those 
leaves and others on the twig, often leaving only the major leaf veins 
intact (Fig. 3). Group A fed on seven of eight adjacent leaves, removing 
43% of the leaf tissue. Group B ate 34% of four adjacent leaves, sub- 
divided and continued feeding on a few more leaves. Four to six days 
after the second feeding bout, the larvae moved up to 3.1 m. During 


188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ines 102 89 65 82 74 69 70 73 65 58 56 55 56 56 55 55 55 54 54 
500 
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Fic. 2. Foraging behavior of tawny emperor caterpillars, from 24 September through 
12 October 1981. EC indicates the position of the egg cluster on the branch. Larval 
numbers per leaf are shown. Arrows indicate periods of larval movement. 


the second migration the larval groups often subdivided, as shown in 
Figs. 1 & 2. At the third set of sites they molted and then fed, staying 
there about five to six days. 

At the first molting site, 72.3 and 71.5% of the original number of 
larvae (Figs. 1 & 2, respectively) were present. About half of the orig- 
inal number of larvae were reaggregated at the second molting site 
(54.6 and 53.9% of those hatching from egg clusters A and B, respec- 
tively). Four factors may account for this larval loss. First, the missing 
larvae may have moved more than 500 cm (length of the major branch- 
es) before the second molt; but this seems unlikely. These larvae use 
silk trails to follow others and remass at new sites. During their migra- 
tion the caterpillars walk along the branches, with leadership contin- 
ually changing as the caterpillars pass others momentarily stopped or 
those backtracking. Consequently, a multi-stranded silk path is depos- 
ited which the last caterpillars follow with less rambling than their 


VOLUME 88, NUMBER 3 189 


es 
DES 


Fic. 8. Feeding site of second-instar tawny emperor caterpillars. Approximately 70% 
of the six distal leaves was eaten. Shaded areas indicate gall tissue that apparently was 
rejected by the caterpillars. 


predecessors. A second factor contributing to larval disappearance may 
have been caterpillars dropping from the trees in response to distur- 
bance. But as early instars, these caterpillars usually drop on silk threads 
and then climb the silk back to their feeding site. A third explanation 
for the decreasing larval numbers may have been that surveying leaves 
during group molting underestimated larval numbers. But the rela- 
tively constant larval numbers during the molting periods suggest that 
few if any larvae were wandering to and from the groups then (see 
“Total/day’for second molts, Figs. 1 & 2). Thus, the fourth factor, 
predation, is probably the major one contributing to larval disappear- 
ance. Only 10 dead larvae were found on the leaves, but predators 
may remove or entirely consume their prey. For instance, pentatomid 
bugs often carry off caterpillars and feed with their prey suspended 
from the beak and off the plant (Evans, 1983). Coccinellid beetle larvae 
may consume all but the head capsule of early instar caterpillars (Stamp, 
pers. observ.). 

The effect of feeding caterpillars aggregating for several days at a 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


site was to concentrate leaf damage. For example, at three feeding 
sites, all of which occurred at the end of branchlets, 79% of the leaf 
tissue of six adjacent leaves, 48% of 19 leaves and 78% of 30 leaves 
were eaten. Consequently, these larval aggregations by way of their 
leaf damage were easy to locate, and presumably predators and para- 
sitoids find them apparent, also. 

These tawny emperor caterpillars exhibited defensive behaviors, such 
as swinging their heads and attempting to bite with their mandibles 
when disturbed. The large, laterally-flattened head capsules with nu- 
merous protuberances on the edge are used in a shield-like manner 
when the caterpillars defend themsleves and may be effective against 
ants and other small predators. These caterpillars are attacked by chal- 
cidoid and ichneumonoid wasps (T. Friedlander, unpubl. data). For 
instance, 68% of 41 larval sites had parasitized caterpillars, indicated 
by the presence of parasitic pupae on or around them (Stamp, unpubl. 
data). (The parasitized caterpillars were left behind when the post- 
molt individuals moved on.) Also, vespid wasps may remove tawny 
emperor caterpillars repeatedly, once they locate an aggregation (T. 
Friedlander, pers. comm.). Thus, it is not clear whether the defenses 
of these caterpillars would be enhanced by aggregation as the defenses 
are for other larvae (e.g., thrashing and regurgitating as with apose- 
matic sawfly larvae, Tostowaryk, 1972). 

The crypticity, movement away from feeding sites before molting, 
and tying leaves at molting sites suggest that tawny emperor caterpil- 
lars may be especially vulnerable to their enemies when molting. These 
caterpillars are less defensive then than during non-molting periods. 
Tawny emperor caterpillars may obtain some advantage from molting 
together, enclosed by leaves bound with silk, or from overwintering 
together (at mid-instar), having tied the deciduous leaves securely with 
multiple silk strands to the tree (Stamp, 1983). Perhaps these advan- 
tages outweigh the possible disadvantages that may arise from group 
feeding (i.e., concentrated leaf damage, which may attract natural 
enemies). 


ACKNOWLEDGMENTS 


I thank Tim Friedlander and Mark Tong for comments on the manuscript. The re- 
search was carried out during a postdoctoral fellowship provided by the Department of 
Zoology at the University of Florida, Gainesville. 


LITERATURE CITED 


EVANS, E. W. 1983. Niche relations of predatory stinkbugs (Podisus spp., Pentatomi- 
dae) attacking tent caterpillars (Malacosoma americanum, Lasiocampidae). Am. 
Mid]. Nat. 109:316-323. 


VOLUME 38, NUMBER 3 19] 


GREENBERG, R. & J. GRADWOHL. 1980. Leaf surface specializations of birds and ar- 
thropods in a Panamanian forest. Oecologia 46:115-124. 

HEINRICH, B. 1979. Foraging strategies of caterpillars: Leaf damage and possible pred- 
ator avoidance strategies. Oecologia 42:325-337. 

& L. COLLINS. 1983. Caterpillar leaf damage, and the game of hide-and-seek 
with birds. Ecology 64:592-602. 

SATO, Y. 1979. Experimental studies on parasitization by Apanteles glomeratus. IV. 
Factors leading a female to the host. Physiol. Entomol. 4:63-70. 

STAMP, N. E. 1988. Overwintering aggregations of hackberry caterpillars (Asterocampa 
clyton: Nymphalidae). J. Lepid. Soc. 37:145. 

TOSTOWARYK, W. 1972. The effect of prey defense on the functional response of 
Podisus modestus (Hemiptera: Pentatomidae) to densities of the sawflies Neodiprion 
swainei and N. pratti banksianae (Hymenoptera: Neodiprionidae). Can. Entomol. 
104:61-69. 





Journal of the Lepidopterists’ Society 
38(3), 1984, 192-193 


HOST SHIFT OF ECPANTHERIA DEFLORATA (ARCTIIDAE) 
FROM AN ANGIOSPERM TO A LIVERWORT 


KEVIN C. SPENCER, LARRY R. HOFFMAN AND DAVID S. SEIGLER 


Department of Plant Biology, University of Illinois, 
Urbana, Illinois 61801 


ABSTRACT. A population of Ecpantheria deflorata Fabricius (Lepidoptera: Arcti- 
idae) was discovered in W. Indiana feeding on Conocephalum conicum (Marchantiales: 
Marchantiaceae), a liverwort. The normal hosts of Ecpantheria in the study area are two 
species of Plantago, and the shift to Conocephalum has occurred despite major differ- 
ences in host plant chemistry. 


Ecpantheria deflorata Fabricius is an arctid moth which has been 
reported to feed on a number of unrelated species in North America 
(Tietz, 1972). These include Brassica oleracea L. (Brassicaceae), He- 
lianthus decapetalus L. (Asteraceae), Robinia pseudocacacia L. (Fa- 
baceae), Euphorbia heterophylla L. and Ricinus communis L. (Eu- 
phorbiaceae), Persea americana Mill. (Lauraceae), Phytolacca ameri- 
cana L. (Phytolaccaceae), Salix sp. (Salicaceae), Viola sp. (Violaceae) 
and Plantago rugellii Dec. and P. lanceolata L. (Plantaginaceae). 

In the Portland Arch Nature Preserve, Fountain Co., Indiana, we 
found that Plantago rugellii and P. lanceolata serve as the major host 
plants for E. deflorata. 

We observed, however, that a number of larvae were grazing exclu- 
sively on Conocephalum conicum L. (Hepaticae), especially in the fall 
of the year. This liverwort forms large mats on sheer rockfaces and is 
very abundant at Portland Arch. Few other lepidopterans are known 
to feed on bryophytes (see Tuskes & Smith, 1984), and not many are 
known from other lower plants (e.g., Euptychia on a lycopsid—Singer 
et al., 1971). Several larvae of various instars were reared on C. coni- 
cum in the laboratory and pupated and emerged normally. 

We consider this host shift to be of interest because of the great 
difference in secondary chemistry between C. conicum and the angio- 
sperm hosts. The chemistry of Conocephalum has been reviewed 
(Markham & Porter, 1978; Spencer, 1979) and the plant has been found 
to contain a large array of mono- and sesquiterpenoids, including up 
to 0.6% dry weight (+) — bornyl acetate, a monoterpene existing in 
opposite chirality to that found in higher plants (Asakawa et al., 1976). 
Some liverwort terpenoids have been shown to inhibit feeding in Lep- 
idoptera (Wada & Munakata, 1971). The major chemical constituents 
of Plantago are iridoid glycosides (Jensen et al., 1975). 

We speculate that some larvae have shifted from Plantago to Co- 


VOLUME 38, NUMBER 3 193 


nocephalum primarily due to the close proximity of infested individ- 
uals of Plantago to Conocephalum covered rocks and to the similar 
texture of the two plants. 

As we have observed a population of Ecpantheria to be present on 
Conocephalum consistently for over 10 years, and given the unique 
chemistry of hepatics which has probably kept lepidopteran herbivores 
in general from utilizing them as host plants, we feel that this host shift 
represents a major event in the population biology of Ecpantheria. 
This may prove to be a useful system for studying population differ- 
entiation across chemical barriers and may represent an incipient spe- 
ciation event. 


ACKNOWLEDGMENTS 


We thank Drs. J. G. Sternburg, G. P. Waldbauer and G. L. Godfrey for confirming 
our insect identification. Vouchers of plant specimens are deposited in the University of 
Illinois Herbarium. 

Thanks are due especially to M. Berenbaum for helpful criticism. 


LITERATURE CITED 


ASAKAWA, Y., M. TOYOTA & T. ARATANI. 1976. (+)-Borny] acetate from Conocepha- 
lum conicum. Proc. Bryol. Soc. Japan 1:155-157. 

JENSEN, S. R., B. J. NIELSEN & R. DAHLGREN. 1975. Iridoid compounds, their occur- 
rence and systematic importance in the angiosperms. Bot. Not. 128:148-180. 

MARKHAM, K. R. & L. J. PORTER. 1978. Chemical constituents of the bryophytes. Pp. 
182-272, in L. Reinhold, J. B. Harborne & T. Swain (eds.). Progress in Phytochem- 
istry, Vol. 5. Pergamon Press, N.Y. 

SINGER, M. C., P. R. EHRLICH & L. E. GILBERT. 1971. Butterfly feeding on lycopsid. 
Science 172:1341-1342. 

SPENCER, K. C. 1979. The chemical constituents of the Hepaticae. Phytochemical 
Bulletin 12:4-19. 

TiETZ, H. M. 1972. An Index to the Described Life Histories, Early Stages and Hosts 
of the Macrolepidoptera of the Continental United States and Canada, Vol. 1. Allyn 
Museum of Entomology, Sarasota, Florida, 1041 p. 

TuskEs, P. M. & N. J. SMITH. 1984. The life history and behavior of Epimartyria 
pardella (Micropterigidae). J. Lepid. Soc. 38:40—46. 

WabDA, K. & K. MunakaTA. 1971. Insect feeding inhibitors in plants. III. Feeding 
inhibitory activity of terpenoids in plants. Agr. Biol. Chem. 35:115-118. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 194-201 


ETHOLOGY OF DEFENSE IN THE APOSEMATIC 
CATERPILLAR PAPILIO MACHAON SYRIACUS 
(PAPILIONIDAE) 


DAviIpD L. EVANS 


Department of Biology, American University of Beirut, 
Beirut, Lebanon 


ABSTRACT. In this investigation I was concerned with two aspects of the defensive 
ensemble of P. machaon syriacus larvae: behaviors which protected them from impend- 
ing predatory attack and population dispersion. There was a comparatively high fre- 
quency of protective behaviors. The high frequency of response may be an adaptation 
against predators which can not recognize the warning signals or those which have a 
way of overcoming the larvae’s defenses. I found that this aposematic insect was not 
commonly in large aggregations. 


Aposematic animals are those which advertise their noxious qualities 
as an anti-predation technique. Clearly, the predators effectively se- 
lecting these aposematic traits will necessarily be able to detect the 
advertisement and gain some advantage in avoiding the noxious prey. 
The predator learns and remembers the undesirability of the prey 
(Evans & Waldbauer, 1982; but see Smith, 1977). An aposematic in- 
dividual may have several different objectionable qualities in its ar- 
mory, each of which may be effective against a different type of pred- 
ator (Edmunds, 1974). 

The larvae of Papilio machaon syriacus Verity (Lepidoptera: Papil- 
ionidae) are brightly colored and fairly obvious at close range. P. ma- 
chaon larvae have been shown to be objectionable to birds (Jarvi et al., 
1981; Wiklund & Jarvi, 1982) and ants (Eisner & Meinwald, 1965). 
This caterpillar seemed to be a good model for investigating certain 
aspects of the aposematic way of life. 

I (1983) had shown that aposematic adult Lepidoptera were less 
likely to perform escape behaviors (elicited by predator-mimicking 
stimuli) than were cryptic, adult Lepidoptera. In this study I wanted 
to determine the frequency of apparently protective behaviors when 
aposematic caterpillars were subjected to various predator-like stimuli 
and the relative rate of habituation with these stimuli. I was also in- 
terested in finding a possible distributional correlate with aposematism. 
Cryptic species generally must maintain low population densities to 
reduce the possibility of search-image formation. Conversely, apose- 
matic animals often form large and conspicuous aggregations (Wiklund 
& Jarvi, 1982). Some aposematic larvae are held at low population 
densities by cannibalism (Williams & Gilbert, 1981). Eruptions of pal- 
atable insects are famous (e.g. locusts, armyworms), on the other hand. 


VOLUME 38, NUMBER 3 195 


METHODS 


I worked in old fields and along roadsides near Jounieh, Lebanon 
from July through September. The last rains generally occur in late 
May. I performed all tests from 1000-1800 h local time when the 
ambient temperature ranged between 30-—40°C. The caterpillars fed 
principally on various above ground portions of Foeniculum vulgare 
Mill. (Umbelliferae). 

I tapped the substrate of resting P. machaon syriacus larvae in order 
to induce a vibration (Evans, 1978) and recorded the response. I per- 
formed this test first, since I found that I often jostled the bushes before 
the end of the tests. Hence, I was more sure that all caterpillars had 
similar treatment. I then applied one of four tactile stimuli: dorsal 
anterior touch (a single tap on the anterior) (group 1, n = 50), anterior 
squeeze (simultaneous bilateral pressure at the anterior) (group 2, n = 
39), dorsal posterior touch (group 8, n = 34), and posterior squeeze 
(group 4, n = 55). I quickly released the bilateral pressure or the tap 
to avoid muting any response. The duration (+0.1 s) and type of re- 
sponse (osmeterial extension, body flexion) were recorded. I then re- 
peated the stimulus and recorded the response type. With dorsal an- 
terior touch, possibly a minimal tactile stimulus, and with posterior 
squeeze, possibly a maximal tactile stimulus, I continued to administer 
the same stimulus every 10 s until the larva either dropped or ceased 
to respond thrice consecutively. This failure to respond three times in 
succession was interpreted as partial evidence of habituation. 

Finally, I changed the second stimulus with a fifth and sixth group 
of larvae. I first administered a dorsal anterior touch and then an 
anterior squeeze to the fifth group (n = 34). With the sixth group (n = 
37), I first applied a posterior dorsal touch then a posterior squeeze. 
The purpose of these last two test series was to compare the reactions 
to a different second stimulus. 

No larva was used in more than one test series. 

I analyzed the data using r X c contingency tables, Poisson analysis, 
and one-way analysis of variance (Snedecor & Cochran, 1980). 


RESULTS 


Initially, I was surprised at how frequently I discovered solitary P. 
machaon syriacus larvae (30.6% were alone). There was a significant 
divergence from the Poisson distribution (P < 0.005) with the majority 
of the high x? value due to the solitaries. Later on, I observed adult 
females ovipositing single eggs ca. 1 m apart. Some large groups (ca. 
60 plants) of F. vulgare had no larvae at all, but some isolated plants 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


were heavily infested (<9 larvae/plant). These multiple infestations 
were quite obvious. The smaller larvae (<15 mm long) were usually 
feeding or resting on umbels where their color patterns were disruptive 
rather than aposematic. Larger caterpillars were rarely on umbels but 
usually on larger stems where they were effectively aposematic. 

The caterpillars reacted to the stimuli by raising the anterior portion 
of the body (illustrated in Eisner & Meinwald, 1965), making a lateral 
thrust with the anterior portion of the body, and/or everting the os- 
meteria. The intensity of these activities varied: 1) In the minimal 
anterior raise, only the head and thoracic legs would rise away from 
the substrate; 2) In the maximum response, the anterior portion of the 
body would be so strongly flexed as to form a “U.” The larva’s lateral 
movement always included the head and thorax, but often the re- 
mainder of the anterior half of the body was also involved. The ever- 
sion of the osmeteria (usually moist) ranged from one-third to fully 
everted. 

When the osmeteria were everted, I was able to smell nothing 44.7% 
of the time. When there was an odor, it was generally similar to butyric 
acid as noted by Eisner and Meinwald (1965). The surprise of the 
osmeterial extension and the odor might induce aversive behavior in a 
potential predator. 

Table 1 illustrates the relative frequencies of behaviors elicited by 
the stimuli when administered initially. The caterpillars were signifi- 
cantly less likely to respond in any obvious way to substrate vibration 
than to the four tactile stimuli (x?, P < 0.001). The elicited responses 
from dorsal anterior touch were not significantly different from those 
with anterior squeeze (x?, P > 0.10). All other frequency comparisons 
were statistically significantly different (x?, P < 0.01). The posterior 
squeeze produced noticeable responses of possibly defensive value in 
96% of the larvae, but substrate vibration elicited an obvious reaction 
in only 20%. Substrate vibration may merely indicate that a leaf glean- 
ing bird or mammal is putting its weight on the stem (Evans, 1978). 
The results show that the posterior squeeze was more likely to stimulate 
a reaction than a dorsal posterior touch. The posterior squeeze approx- 
imates a grasp by a bird’s beak and so is more similar to a real threat. 

The mean durations of the various behaviors are also noted in Table 
1. The means were not statistically significantly different (ANOVA, 
P > 0.05). 

I wished to determine whether the larvae normally repeated the 
same behavior after receiving a seond similar stimulus. Fig. 1 illustrates 
the frequency of behaviors with the group | caterpillars as an example. 
Forty-one larvae exhibited similar behavior after the second anterior 
dorsal touch; only nine had different responses the second time. This 


197 


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198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Response after first Response after second 
anterior dorsal touch anterior dorsal touch 
no response no response (23) 






Osmeteria extended (II) 





Osmeteria extended 





(I) 

Lateral 
(1) 
a 


VOLUME 38, NUMBER 3 199 


general pattern of behavior occurred with the other three tactile stim- 
uli when I repeated each of them. No set of second responses in any 
group was statistically significantly different from the set of first re- 
sponses in that group (x?, P = 0.10). 

In groups 5 and 6, I applied a different second tactile stimulus. The 
response frequency evoked by the anterior squeeze (as a second stim- 
ulus) was not significantly different from that appearing after the sec- 
ond anterior touch (x?, P > 0.10). However, the frequency was signif- 
icantly different when the second stimulus was a posterior squeeze 
compared to when it was a repeated posterior touch (x?, P < 0.05). 

In the habituation test, I found that the larvae continuing to receive 
the anterior dorsal touch exhibited some type of response slightly fewer 
times (x = 20.0 + 16.87) than with the posterior squeeze (x = 20.5 + 
13.75). Three of the latter group eventually dropped but none of the 
former. The eventual failure to respond was probably not due to fa- 
tigue, since several of the non-responding larvae crawled away after I 
stopped applying the stimuli. 


DISCUSSION 


The highly localized groupings of larval P. machaon syriacus added 
to the overall impression of conspicuousness. Aposematic caterpillers 
often seem to feed in obvious locations (Heinrich, 1979). These larvae 
are distasteful to avian insectivores, and the caterpillars usually survive 
an attack from birds (Jarvi et al., 1981; Wiklund & Jarvi, 1982). The 
numerous aposematic larvae may act as a supernormal releaser in stim- 
ulating aversive behavior in the predator (Cott, 1940). Individual fit- 
ness may be increased in large groups of aposematic larvae since para- 
sitoid-related mortality is reduced (Baker, 1970). Therefore, the high 
incidence of solitary individuals in this warningly colored species is 
surprising. 

The degree of responsiveness to the tactile stimuli is also surprising 
in light of earlier work (Evans, 1983). The relatively high frequencies 
of responses and the reduced gregariousness could be rationalized if a 
large component of the mortality of the larvae were due to ants or 


—_— 


Fic. 1. Frequency of reactions after a first dorsal anterior touch and then a second 
dorsal anterior touch. The width of the arrows is roughly proportional to the number of 
individuals performing the second act. O.E. = osmeteria extended; R.A. = raised ante- 
rior; N.R. = no observable response; L.A.M. = lateral anterior movement. Some activities 
are performed simultaneously. The parentheses at the right show the actual number 
performing the action. n = 50. 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


some other predator where learning plays a minor role in prey selection 
or where there would be little innate recognition of noxious prey. The 
most frequent responses (Table 1) included osmeterial extension as at 
least one component. The osmeterial secretions act primarily against 
ant predation (Eisner & Meinwald, 1965) but have little, if any, role 
in defense against bird predation (Jarvi et al., 1981). Eisner and Mein- 
wald (1965) note, however, that ants can exhaust the osmeterial secre- 
tions of these larvae by making repeated attacks. Whether or not such 
attacks occur in nature is unreported. The larva in such a situation 
might survive by throwing ants off with vigorous body thrusts. 

The behavioral responses seem to be modal or fixed action patterns 
to the extent that they were stereotyped and appeared to have little 
learned component. The patterns of behavior were fixed since the same 
response was most often given to the same stimulus the second time. 
Most of the larvae eventually ceased to respond defensively. It appears 
that there was habituation. 


CONCLUSIONS 


The aposematic defensive ensemble implies a higher relative thresh- 
old for release of active protective behaviors (Evans, 1983). This prin- 
ciple is contingent upon a predator recognizing the aposematic signal 
and then avoiding contact with the noxious item. The results of this 
study suggest that some predators do not recognize the aposematic 
signal and are consistently warded off only by repeated active defenses. 


ACKNOWLEDGMENTS 


I wish to thank Henreitte Khouweiry and the Anton Sfeir family for their assistance 
while I was in Jounieh, Lebanon. A grant from the Faculty of Arts and Sciences of the 
American University of Beirut supported this research. Drs. L. Young and L. Squires 
offered helpful suggestions in the preparation of this article. Dr. Samir Deeb deserves a 
special note of thanks for all that he did last summer. Amin Abou-Samra produced the 
figure. 


LITERATURE CITED 


BAKER, R. R. 1970. Bird predation as a selective pressure on the immature stages of 
the cabbage butterflies, Pieris rapae and P. brassicae. J. Zool., Lond. 162:43-59. 
Cott, H. B. 1940. Adaptive Coloration in Animals. Methuen & Co., London. 508 pp. 
EDMUNDS, M. 1974. Defence in Animals. Longman Group Limited, Harlow, Essex, 
United Kingdom. 357 pp. 

EISNER, T. & Y. C. MEINWALD. 1965. Defensive secretion of a caterpillar (Papilio). 
Science 150:1733-1735. 

Evans, D. L. 1978. Defensive behavior in Callosamia promethea and Hyalophora 
cecropia (Lepidoptera: Saturniidae). Am. Midl. Nat. 100:475—479. 

1983. Relative defensive behaviour of some moths and the implications to 

predator-prey interactions. Entomol. Exp. Appl. 33:103-111. 

& G. P. WALDBAUER. 1982. Behavior of adult and naive birds when presented 

with a bumblebee and its mimic. Z. Tierpsychol. 59:247-260. 








VOLUME 38, NUMBER 3 201 


HEINRICH, B. 1979. Foraging strategies of caterpillars, leaf damage, and possible pred- 
ator avoidance strategies. Oecologia 42:325-337. 

JARVI, T., B. SILLEN-TULLBERG & C. W. KLUND. 1981. The cost of being aposematic. 
An experimental study of predation on larvae of Papilio machaon by the great tit, 
Parus major. Oikos 36:267-272. 

SMITH, S. M. 1977. Coral-snake pattern recognition and stimulus generalization by 
naive great kiskadees (Aves: Tyrannidae). Nature 265:535-536. 

SNEDECOR, G. W. & W. G. COCHRAN. 1980. Statistical Methods, 7th ed. Iowa State 
University Press, Ames, Iowa. 507 pp. 

WIKLUND, C. & T. JARvI. 1982. Survival of distasteful insects after being attacked by 
naive birds: A reappraisal of the theory of aposematic coloration evolving through 
individual selection. Evolution 36:998-1002. 

WILLIAMS, K. S. & L. E. GILBERT. 1981. Insects as selective agents on vegetative 
morphology: Egg mimicry reduces egg laying by butterflies. Science 212:467—469. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 202-208 


THE EGG OF HOFMANNOPHILA PSEUDOSPRETELLA 
(OECOPHORIDAE): FINE STRUCTURE OF THE CHORION! 


RICHARD T. ARBOGAST AND RICHARD VAN BYRD 


Stored-Product Insects Research and Development Laboratory, 
Agricultural Research Service, USDA, Savannah, Georgia 31403 


GEORGES CHAUVIN 


Laboratoire de Biologie Animale, Université de Rennes, 
35042 Rennes Cedex, France 


RUDOLPH G. STRONG 


Department of Entomology, University of California, 
Riverside, California 92521 


ABSTRACT. The egg of Hofmannophila pseudospretella (Stainton) was studied by 
scanning and transmission electron microscopy. The egg is usually obovoid but varies to 
ellipsoid or subcylindrical (0.58 x 0.41 mm). The basic pattern of sculpturing consists of 
low-lying longitudinal ridges joined by indistinct transverse ridges with the ridge inter- 
sections slightly elevated. This pattern is sometimes poorly developed, but the slight 
prominences formed by intersecting ridges are always evident. The surface of the chorion 
has a wrinkled or granular texture. There are 3 to 5 micropylar canals opening into an 
anterior pit which is surrounded by a rosette of rather short, petal-shaped primary cells. 
The primary cells are in turn partially or completely surrounded by series of secondary 
and tertiary cells. The aeropyles open on slight prominences near the anterior and pos- 
terior ends of the egg. Openings are quite abundant in these areas, but there are none 
elsewhere. Typically, the openings are funnel-shaped and may or may not be surrounded 
by collars. The chorion averages 4.23 um thick, and in general structure is similar to that 
of other lepidopteran eggs. 


The brown house moth, Hofmannophila pseudospretella (Stainton), 
is a cosmopolitan, household, mill, and storage pest. The larvae are 
omnivorous scavengers and attack a wide range of plant and animal 
products. In North America, H. pseudospretella is found from Cali- 
fornia north to British Columbia and east to Manitoba; there are iso- 
lated eastern records from Pennsylvania and southwest Greenland 
(Hodges, 1974). It has been recorded from cantaloupe seed, celery seed, 
fish meal, grain, mixed feed, lima beans, and milo in California (Strong 
& Okumura, 1958; Okumura & Strong, 1965). In Europe, H. pseudo- 
spretella is a common pest of stored products. Woodroffe (1951) re- 
ported that it is widely distributed in Britain, where it occurs in dwell- 
ings, stores, and mills as well as outdoors in bird nests. It occasionally 
becomes a major pest attacking bulk wheat, bagged flour, and other 
stored commodities. In the home, it is most often important as a clothes 


‘Mention of a proprietary product in this paper does not constitute an endorsement of the product by the USDA. 


VOLUME 38, NUMBER 3 203 


moth. It is common and generally distributed in New Zealand where 
it is most often a pest of carpets (Somerfield, 1981). 

Woodroffe (1951) gave an account of the life-history of H. pseu- 
dospretella, including a brief description of the egg. However, there 
have been no detailed studies of its chorionic structure. The study 
reported here was conducted as part of a project to characterize the 
eggs of stored-product insects and to facilitate their identification. The 
only other oecophorid egg that has been studied is that of the white- 
shouldered house moth, Endrosis sarcitrella (L.) (Arbogast et al., 1983). 


MATERIALS AND METHODS 


Laboratory cultures of H. pseudospretella were established with 
adults that emerged from food packets (Strong, 1970) placed in an old 
shed and an old barn near Castroville, California. Voucher specimens 
have been deposited in the U.S. National Museum of Natural History. 
The moths were reared in the laboratory at 25 + 1°C on 950 ml 
quantities of cracked wheat in 3.79 | jars. Cultures were maintained 
in a room at 60 + 5% RH, but additional moisture was provided by a 
watering device in each jar. This consisted of a plastic vial (6 cm 
deep X 4 cm inside diameter) filled with water and covered with a 
piece of 11 cm filter paper that served as a wick. The filter paper was 
held in place by the vial’s snap-on cap, which had a hole 25 mm in 
diameter cut in its center. The vial was placed in an inverted position 
on the bottom of the jar, and wheat was poured on top of it. 

Eggs were collected by confining moths in 0.95 | jars without food. 
Each jar was covered with screen secured by a screw-type lid, and 
each contained a piece of pleated black construction paper to provide 
a resting place. Jars containing moths were held in a desiccator over a 
saturated solution of KNO,, which provided an RH of ca. 92%. The 
moths oviposited freely in this situation, and the eggs, which adhered 
very lightly or not at all to the surfaces on which they were deposited, 
were collected by shaking them onto a piece of paper. After they were 
collected, the eggs were washed by gentle agitation or sonication for 
ca. 5 min in a 1% solution of Triton X-100®, rinsed in distilled water 
and air dried. 

For examination in the scanning electron microscope (SEM), the 
eggs were mounted with double-sided tape on SEM stubs and sputter- 
coated with gold. They were examined with an International Scientific 
Instruments, M-7® SEM at 15 kV. Approximately 250 eggs selected 
arbitrarily from groups laid by ca. 30 females were examined. Length 
and width were determined from a sample of 33 eggs. Measurements 
were made on the display screen of the microscope at a magnification 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of x150. The diameter of 18 aeropylar openings was determined from 
micrographs at either <15,000 or x20,000. The openings measured 
were on 13 different eggs. Counts of primary cells were made on a 
sample of 20 eggs, either from the screen or from micrographs. All 
measurements and counts are given as mean + standard deviation 
(S.D.). The terminology used in describing the structural features of 
the chorion is the same as used by Arbogast et al. (1980). 

For examination by transmission electron microscope, the eggs were 
punctured with a minuten pin, fixed overnight in cold glutaraldehyde 
(5% in Millonig’s buffer, pH 7.4), rinsed in Millonig’s buffer, and post- 
fixed overnight in osmium tetroxide (1% in Millonig’s buffer, pH 7.4). 
After fixation, the eggs were rinsed in Millonig’s buffer and dehydrated 
in a graded series of water-ethanol solutions followed by ethanol and 
propylene oxide. Initial infiltration in a 1:1 mixture of propylene oxide 
and embedding resin (Araldite 6005®) overnight was followed by in- 
filtration overnight in a 1:2 mixture of propylene oxide and resin and 
infiltration for three days in pure resin. After infiltration, the eggs were 
transferred to resin which was then cured at 48°C overnight. Sections 
were cut using a glass knife on a Porter Blum MT-2B® ultramicrotome 
and stained by flotation of grids on a 1% solution of uranyl acetate in 
water for 7.5 min followed by flotation on Reynold’s lead citrate for 
2.5 min. The sections were examined in a Phillips EM-200®. The thick- 
ness of the chorion and each chorionic layer is given as mean + S.D. 
All means are based on measurements of 14 sections taken from a total 
of four eggs that were selected arbitrarily from a group laid by 36 
females. 

Eggs from adults collected in France, near Rennes, were also studied 
by the same methods. 


RESULTS AND DISCUSSION 


We found no differences between eggs of California H. pseudo- 
spretella and those collected in France. The eggs is usually obovoid 
but varies to ellipsoid or subcylindrical, 0.58 + 0.04 mm long x 0.41 
+ 0.02 mm in diameter at its broadest point (Fig. 1). Woodroffe (1951) 
described the egg as hard and shiny, oval and tapering toward one 
end. He noted that the eggs vary considerably in size and color and 
was able to distinguish two extreme types: (1) small and white, aver- 
aging 0.490 mm long and (2) large and yellow, averaging 0.595 mm 
long. Although Woodroffe stated that these differences persist through- 
out the incubation period, we were unable to distinguish such distinct 
types among the eggs we examined. 

The egg is not boldly marked. Its basic pattern of sculpturing consists 
of low-lying longitudinal ridges joined by indistinct transverse ridges, 


VOLUME 88, NUMBER 3 205 





st aaa 


Fics. 1-4. Egg of Hofmannophila pseudospretella. 1, Lateral view of whole egg, 
anterior pole on left (x80). 2, Anterior end showing micropylar area and aeropyles 
(arrows) (570). 3, Micropylar area showing central pit (arrow) with four micropylar 
canals opening on its periphery and rosette of cells surrounding the pit (x1140). 4, 
Aeropyle near posterior end of egg ( x 8590); the inner layer of chorion, which forms the 
floor of the trabecular layer, is visible at the bottom of the opening. 


the junctures being slightly elevated. This pattern is sometimes poorly 
developed and the ridges almost imperceptible, but the slight promi- 
nences formed by intersecting ridges are always evident, especially 
near the poles of the egg. The surface of the chorion has a wrinkled 
or granular texture (Figs. 2 & 8). 

There are 3 to 5 micropylar canals opening into a central micropylar 
pit at the anterior pole of the egg (Figs. 2 & 3). The pit is surrounded 
by a rosette of 5-8 (6.9 + 0.9) rather short, petal-shaped primary cells, 
which are in turn partially or completely surrounded by series of sec- 
ondary and tertiary cells. The primary cells, and usually the secondary 
cells as well, are outlined by prominent carinae, and often carinal spurs 
extend into the cell discs. The tertiary cells, on the other hand, are 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 5. Egg of Hofmannophila pseudospretella: thin section through the chorion and 
underlying membranes. EL, external layer of chorion; IL, internal layer of chorion; PL, 
principal layer of chorion; TL, trabecular layer of chorion; VM, vitelline membrane; Y, 
yolk material. (<18,340). 


often outlined only by low-lying ridges and are thus poorly defined 
(Fig. 2). 

The aeropyles open on slight prominences at the anterior and pos- 
terior ends of the egg (Fig. 2). Openings are relatively abundant in 
these areas but are absent elsewhere. Typically, the openings are fun- 
nel-shaped and may be surrounded by collars (Figs. 2 & 4). The di- 
ameter at the narrow end of the funnel ranges from less than 0.5 to 
more than 2.5 wm (1.0 + 0.6 wm). 


VOLUME 38, NUMBER 3 207 


In sections (Fig. 5), the surface of the chorion appears quite smooth 
and devoid of mucilaginous colleterial secretions, which accounts for 
the failure of the eggs to adhere firmly to the substrate on which they 
are laid. There is no furrowing as in E. sarcitrella (Arbogast et al., 
1988). The chorion averages 4.23 + 0.76 um in thickness, much thicker 
than the chorion of E. sarcitrella, which ranges from ca. 0.6 to 1.1 wm. 
The chorion of H. pseudospretella consists of four distinct layers (Fig. 
5): an external layer (EL) (0.07 + 0.03 um), a lamellate principal layer 
(PL) (3.90 + 0.72 um), a trabecular layer (TL) (0.21 + 0.05 um) which 
consists of air held between vertical columns (trabeculae) and com- 
prises the intrachorionic respiratory meshwork of the eggs, and an 
inner layer (IL) (0.06 + 0.02 um). Two layers can be distinguished 
within the principal layer by the orientation of the lamellae. These are 
parallel to the surface in the inner % of the principal layer but are 
usually oblique in the outer %. This orientation as well as thickness 
may account for the resilience of the eggshell noted by Woodroffe 
(1951), who stated that “if an attempt is made to crush ... [an egg] 
with a dissecting needle, . . . [it] will usually spring undamaged from 
beneath the needle.” In the section figured, the vitelline membrane 
(VM) and yolk material (Y) are visible beneath the chorion. The egg 
from which this section was taken was newly laid; the vitelline mem- 
brane has not yet condensed, and the serosa and serosal cuticle have 
not developed. 

The eggs of H. pseudospretella and E. sarcitrella resemble some- 
what the eggs of Tineidae, but tineid eggs can be distinguished by the 
presence of microperforations in the surface of their chorion (Arbogast 
et al., 1980; Chauvin, 1977). The eggs of H. pseudospretella and E. 
sarcitrella have well-defined primary and secondary cells, and some- 
times tertiary cells, in the micropylar area, but the remainder of the 
surface lacks cells, or has cells that are at best only faintly outlined and 
barely discernible. The eggs of all other stored-product moths that have 
been studied (except Tineidae) are marked by extensive reticulate pat- 
terns of well-defined cells and ridges; usually, these cover the entire 
surface of the egg (Arbogast et al., 1980; Arbogast & Byrd, 1981). Eggs 
of E. sarcitrella can easily be distinguished from those of H. pseudo- 
spretella by the maze-like pattern of closely spaced sinuous ridges that 
covers their entire surface and by the absence of aeropyles near their 
poles (Arbogast et al., 1983). In general structure, the chorion of H. 
pseudospretella is similar to most other lepidopteran eggs that have 
been studied (see, for example, Barbier & Chauvin, 1974; Chauvin & 
Barbier, 1976; Chauvin et al., 1974; Salkeld, 1973). The chorion is 
about the same thickness as that of Galleria mellonella (L.) (8.1 to 4.2 


208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mm) (Barbier and Chauvin, 1974) but is thicker than that of Tinea 
pellionella L. (2.5 to 3.5 um) or Tineola bisselliella (Hummel) (0.5 to 
1.0 wm) (Chauvin, 1977). 


ACKNOWLEDGMENT 


We are indebted to Margaret Carthon, Biological Technician at the Stored-Product 
Insects Research and Development Laboratory for rearing the moths and for assistance 
in collecting the eggs. 


LITERATURE CITED 


ARBOGAST, R. T. & R. V. ByrD. 1981. External morphology of the eggs of the meal 
moth, Pyralis farinalis (L.), and the murky meal moth, Aglossa caprealis (Hiibner) 
(Lepidoptera: Pyralidae). Int. J. Insect Morphol. Embryol. 10:325-329. 

, G. CHAUVIN, R. G. STRONG & R. V. BYRD. 1988. The egg of Endrosis sarci- 

trella (Lepidoptera: Oecophoridae): Fine structure of the chorion. J. Stored Prod. 

Res. 19:63-68. 

, G. L. LEcaTo & R. V. ByrD. 1980. External morphology of some eggs of 
stored-product moths (Lepidoptera: Pyralidae, Gelechiidae, Tineidae). Int. J. Insect 
Morphol. Embryol. 9:165-177. 

BARBIER, R. & G. CHAUVIN. 1974. Ultrastructure et réle des aéropyles et des enveloppes 
de l’oeuf de Galleria mellonella. J. Insect. Physiol. 20:809-820. 

CHAUVIN, G. 1977. Contribution a l'étude des insectes kératophages (Lepidoptera, 
Tineidae): Leur principales adaptations a la vie en milieu sec. Thesis, Université de 
Rennes. 295 pp. 

& R. BARBIER. 1976. Développement des oeufs en fonction de lhumidité et 

structure de leurs enveloppes chez quatre lépidoptéres Tineidae: Monopis rusticella 

Clerck, Trichophaga tapetzella L., Tineola bisselliella Hum. et Tinea pellionella 

L. Act. 97° Congres Natn. Soc. Savantes 3:627-643. 

, R. RAHN & R. BARBIER. 1974. Comparaison des oeufs des lépidoptéres Phalera 
bucephala L. (Ceruridae), Acrolepia assectella Z. et Plutella maculipennia Cutt. 
(Plutellidae): Morphologie et ultrastructures particuliéres du chorion au contact du 
support végétal. Int. J. Insect Morphol. Embryol. 3:247-256. 

HopcEs, R. W. 1974. Gelechioidea: Oecophoridae (in part). Fasicle 6.2 (142 pp.) in 
R. B. Dominick et al. (eds.). Moths of America North of Mexico. E. W. Classey Ltd., 
London. 

OxkuMuRA, G. T. & R. G. STRONG. 1965. Insects and mites associated with stored food 
and seeds in California. Part II. Bull. Dep. Agric. Calif. 54:13-23. 

SALKELD, E. H. 1973. The chorionic architecture and shell structure of Amathes c-ni- 
grum (Lepidoptera: Noctuidae). Can. Entomol. 105:1-10. 

SOMERFIELD, K.G. 1981. Recent aspects of stored product entomology in New Zealand. 
N.Z. J. Agr. Res. 24:403-408. 

STRONG, R. G. 1970. Distribution and relative abundance of stored-product insects in 
California: A method of obtaining sample populations. J. Econ. Entomol. 63:591- 
596. 














& G. T. OKUMURA. 1958. Insects and mites associated with stored foods and 
seeds in California. Bull. Dep. Agric. Calif. 47:233-249. 

WooproFFE, G. E. 1951. A life-history study of the brown house moth, Hofmanno- 
phila pseudospretella (Staint.) (Lep., Oecophoridae). Bull. Entomol. Res. 41:529- 
508. 





Journal of the Lepidopterists’ Society 
38(3), 1984, 209-219 


THE DYNAMICS OF ADULT DANAUS PLEXIPPUS L. 
(DANAIDAE) WITHIN PATCHES OF ITS FOOD PLANT, 
ASCLEPIAS SPP. 


M. P. ZALUCKI 


Department of Entomology, University of Queensland, 
St. Lucia, Queensland 4067, Australia 


R. L. KITCHING 


School of Australian Environmental Studies, Griffith University, 
Nathan, Queensland 4111, Australia 


ABSTRACT. Mark-recapture studies of adults of Danaus plexippus at four sites in 
southeast Queensland have been made over a ten month period. Despite unavoidable 
difficulties associated with non-compliance with the basic assumptions of standard mark- 
recapture techniques, population trends have been identified, abundance levels having 
late summer and late autumn peaks, declining to low levels in winter. Longevity of 
individuals, estimated as the “mean minimum life-span,” was 12.4 days overall (13.5 for 
males, 10.0 for females). These estimates are regarded as conservative. Males predomi- 
nated in all four sites but a seasonal pattern was evident with the proportion of males 
declining to below 50% at the end of summer. In the remainder of the sampling period 
the proportion of males varied from 50 to 75%. Separate estimates of population turnover 
in a single patch were obtained—within four days, an estimated 50% of the local pop- 
ulation had been replaced by “new” individuals about half of which were newly emerged 
and half, presumably, immigrants. 

The male-biased sex ratios are hypothesized to be due, at least in part, to aggressive 
male-female interactions with resident males, reducing within-patch residence times of 
females. The overall vagility of the species and the difficulty of studying events occurring 
between patches makes interpretation of the adult dynamics in this species difficult. 
Further work, investigating the role of intersexual behavior in determining within patch 
dynamics is in progress. 


The distribution of host plants has been shown to influence the dis- 
persion, abundance and survival of both larval and adult butterflies 
(Dethier, 1959; Ehrlich, 1965; Singer, 1971; Sharp & Parks, 1978; Za- 
lucki, 1981a). Monarch butterflies (Danaus plexippus L.) in Australia 
lay eggs on introduced Asclepias spp. (milkweeds). These plants often 
grow in large dense patches (Zalucki et al., 1981). In this study we use 
mark-recapture techniques to investigate abundance, longevity and 
movement of adult monarchs at and between four milkweed patches 
in southeast Queensland. The monarch is native to North America and 
was first noted in Australia around 1871. In southeast Queensland the 
species breeds throughout the year (Smithers, 1977). 


MATERIALS AND METHODS 


The four study sites were located southwest of Brisbane (centering 
on 27°28'S; 158°1'E). The sites were widely spaced (5-16 km apart, 
Fig. 1) and there were numerous patches of milkweed in the interven- 
ing areas (see Zalucki et al., 1981 for a description of patch dispersion). 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Logan 
Village 





Tamborine 


Fic. 1. Map showing location of study sites 1, 2, 3 and 4 (A), major roads (—) and 
urban centers. (--) indicates direction of movement and, the number of arrowheads, those 
butterflies marked at one site and recaptured at another. 


Each site was sampled by 30-40 minutes netting once a week. Site 
1 was sampled from 8 February to 5 July 1978 and site 3 from 2 
February to 28 June 1978, whereas, sites 2 and 4 were sampled from 
2 February to 22 November 1978 inclusive. Butterflies were tagged 
using the alar method outlined by Urquhart (1960), sexed and classified 
as either fresh (wings soft), old (wings chipped, color very faded) or 
middle aged (wings slightly chipped, little fading). 

Population size, rates of recruitment and losses from a site were 


VOLUME 38, NUMBER 3 DA | 


TABLE 1. Summary of mark-recapture observations by site and sex. 


Males Females 
Site Marks Recapt. % recapt. Marks Recapt. % recapt. % males 
1 228 18 U8) 142 6 4.2 O1G27% 
2 249 32 12.9 143 23 16.1 0.64** 
3 237 22 9.3 132 8 6.2 0.64** 
4 bee es 19.8 422 57 13.5 0.58** 
1299 185 14.2 839 94 2 OG 144 


** Significantly different from 0.5, P < 0.05. 


estimated from the recaptures using the Jolly-Seber method (Jolly, 1965; 
Southwood, 1966; Seber, 1973). Longevity of marked individuals was 
estimated as the “mean minimum life-span”’ (i.e. sum of days survived 
by all recaptures/total recaptured, see Ehrlich & Gilbert, 1973). The 
recapture at one site of individuals marked at another provided crude 
estimates of potential cross-country movements. The sex ratio of the 
catch was also recorded at each site on each sampling occasion. 

In addition on one occasion, virtually all butterflies at site 4 were 
captured and marked on one day. The purpose of this exercise was 
threefold: (1) to determine whether the sample sex ratios were accu- 
rate; (2) to check population size estimates; and, (3) to check the rate 
at which a marked population is diluted by unmarked insects. 


RESULTS 


During the study period a total of 2138 butterflies were netted at 
the four sites, and of these, 279 (13%) were recaptured at least once; 
with the exception of site 2, a higher percentage of males was recap- 
tured than females (Table 1). 

The populations at the four sites were not isolated from each other. 
Five marked individuals were recaptured at a site other than the one 
at which they were marked (recapture distances 6.2, 6.2, 6.2, 7.8, 15.6 
km) and one specimen was captured 0.7 km from site 4 (Fig. 1). From 
these interpatch recaptures we estimated that cross-country moves can 
proceed at a rate of ca. 1.5 km/day (range 0.7-2.48). 


Population Estimates 


Due to the low recapture rate at each site each week, male and 
female recaptures were combined to give weekly population estimates 
for each site (Fig. 2). The estimated population sizes, dilution and loss 
rates at each patch were characterized by their large variability. Field 
experience indicated that the population at each site did not fluctuate 
to the extent that the estimates suggested. The calculated fluctuations 


212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


600 


> 500 


( 


a—--« Site 4 


Pi 


BSS 
fo) 
(eo) 


ol 
° 
(o) 


N) 
{e) 
fe) 


| 
i} 
! 
! 
| 
i 
i 
\ 


Estimated Population Size ( 





100 Me 
\ i Le ; 
0 
O Feb Mar Apr May Jun Jul Aug Sep Oct Nov 
Summer Autumn Winter Spring 
MONTH 


Fic. 2. Estimates of population sizes (P, given by combining male and female recap- 
tures) for each site. 


were due to the failure of some of the assumptions of the method of 
estimation and the low rates of recapture. 

One assumption is that recaptures be random. Marked females were 
recaptured randomly (z = 0.38588, P > 0.5); whereas, males were not 
(z = 2.3294, P < 0.05). The other assumption that is violated is that 
migration be permanent. Direct evidence for this occurred during the 
patch clearing exercise. Two tagged specimens where recaptured at 
site 4 before and after but not during the patch clearing exercise (see 
below). 

Despite these problems, population trends over the sampling period 
are apparent (Fig. 2). Populations are high in late summer (100-200 
butterflies per patch in February, Fig. 2), declining in early autumn 
(to around 50/patch in March), increasing again in late autumn to 
about 100/patch (April, May) and declining to low levels (10-50/patch) 
in winter. Populations increase in spring to achieve the high levels 
observed in summer. 

Due to the variability of the various estimates obtained using the 
Jolly-Seber methods, the associated estimates of survivorship are un- 
reliable. For these reasons we have analyzed longevity (survivorship) 
using a different method. 


VOLUME 38, NUMBER 3 213 


TABLE 2. Longevity? of marked D. plexippus. 


This study North America? Australiac 
Min. July— Min. 
Min. days days Aug 16¢ All days 
surviva $ g Ap seal Summer months T survival $+2 
0-5 4 33 37 3.5 720 
7 93 67 160 5-10 3) 20 25 10.5 360 
10-15 ih 13 20 17.5 214 
14 47 20 +67 15-20 4 6 10 24.5 133 
20-25 4 9 31.5 76 
21 23 5 28 25-30 2 4 6 38.5 57 
30-35 4 4 45.5 53 
28 16 i 17 35-40 3 3 02.9 52 
40-45 iI 3 4 09.9 De, 
35 4 4 45-50 ] L 66.5 20 
50-55 4 4 73.5 15 
49 1 1 50-60 80.5 9 
60-65 2 2 87.5 17 
56 1 1 65-70 if 1 94.5 rh 
70-75 101.5 8 
70 1 il 75-80 108.5 ia 
>80 6 6 >115.5 85 
Total 185 945219 Total 27 105 132 1806 
Mean min. 
life-span 
(days) HSto / LO 12.4 14.5 2Owre 2082 19:9 


* Based on Ehrlich and Gilbert (1973). [(No. known age x days since capture)/Total recaptures] = Mean minimum 
ife-span. 
> Taken from Urquhart (1960, table II, p. 291). 
© Calculations based on the midpoint of the interval. Those recaptured on the same day have been ignored. 
4 Marked and recaptured during summer months. 
© Taken from Smithers (1973, table II). 


Longevity 


The “mean minimum life-span” for all individuals is about 12.4 
days; 13.5 days for males and 10 days for females (Table 2). These 
values greatly underestimate longevity, reflecting as they do, the time 
spent in a patch (see below). This is determined by movements into 
and out of patches rather than by permanent removal due to death. 
Our estimates of life-span are similar to those calculated for the sum- 
mer breeding population of D. plexippus in North America (Table 2). 
This presumably reflects a similar underlying population structure. The 
estimated life-spans are consequently less than the 40 days recorded 
for adults in captivity (Urquhart, 1960; Munger & Harriss, 1970; Za- 
lucki, 1981b) and of adults marked and recaptured over the years in 
North America (Table 2). The estimated longevities based on Smithers 
(1978, table II) are also longer than our estimates. Smithers’ recaptures 
are based mainly on butterflies from around Sydney, N.S.W., where 


214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Mean minimum life-span (in days) estimates* for sites 1-4 by sex and age. 


Males Females 
Site Fresh> Other* Fresh? Other 
1] 8.8 11.5 d 8.2 
2 9.4 14.9 7.0 10.1 
3 9.5 10.0 q 11.4 
4 15.9 NS 7 7.0 10.3 
Pooled estimate 13.0 13.7 7.0 9.9 


* Formula as in Table 4. 

> Newly emerged, wings soft. 

© Not newly emerged. 

4 None caught in this category. 


adults hibernate. That adults lived longer than indicated by recapture 
data is evidenced by the good condition of most adults (even the long- 
est-lived) when they were last captured compared with their condition 
at the time of first capture. 

The mean minimum life spans were calculated for each patch site 
with individuals categorized by age on the basis of wing wear (Table 
3). These estimates are an indication of residence time in a patch. 
Females, both fresh and otherwise, have lower residence times than 
males (Table 3). We interpret this as being because females are more 
likely to leave a patch (i.e., have a shorter residence time) than males. 


Sex Ratios 


Males predominate at all patch sites (Table 1) and show similar 
changes in relative abundance over the study period across all four 
sites (Fig. 3; Spearman rank correlation coefficients: S1 <x S3, 0.367, 
0.01 < P < 0.05; S1 x S4, 0.224, P > 0.005; S838 x S4, 0.728, P < 0.001; 
site 2 was ignored due to low sample sizes as were all samples with 
<10 individuals). All sites show a change from significantly more than 
50% males, to significantly less than 50% males around early autumn 
(March, April). At other times of the year the sex ratio fluctuates from 
75% to 50% males (Fig. 3). These changes may be due to sampling 
bias, changes in the sex ratio at birth, changes in male/female migra- 
tion rates or male/female death rates. 

To test if samples were biased we compared the estimates of sex 
ratio before and after the patch clearing exercise. This exercise gave 
the actual sex ratio in the patch on 8 July as 61% males. This does not 
differ from the sex ratio estimated by samples taken before (66%) and 
after this date (69%). On the strength of this result we feel that the 
sample estimates of the sex ratio probably indicate real fluctuations in 
the percentage of males. These fluctuations are not due to the produc- 


VOLUME 38, NUMBER 3 DAs) 


Percent Males 





Fics. 3a—d. Percentages of males for each site plotted against time: site 1 (a), site 2 
(b), site 3 (e) and site 4 (d). Solid symbols indicate significant deviation from 50% (P < 
0.01). Open symbols denote non-significance. Double symbols indicate sample size small 
(<10). — average percent males across all sites shown in (d). 


tion of all male broods—the sex ratio of newly emerged butterflies 
pooled over time for each site and the changes in the sex ratio at birth 
over time (pooled across sites) do not differ significantly from 50/50. 
This agrees with the sex ratio of laboratory reared adults (Zalucki, 
1981c). 


Population Turnover 


Since, as far as we could tell, every adult was marked at site 4 on 8 
July, a sample taken on the 12th should reveal what fraction had left, 
how many migrants had arrived and how many had emerged from 
around the patch (Table 4). Assuming that the population was about 
the same size on the 12th as on the 8th, then the sample taken on the 
12th represents about half the population. On this basis multiplying 
the sample values on the 12th by two gives an estimate of the various 
marked and unmarked fractions in the population on that date. 

On the basis of these calculations we suggest that about half the 
individuals marked on the 8th had left, and this was similar to the 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Mark-recapture history of adults netted on 8 and 12 July. 


Date of first capture for marked insects 


Total Un- ee es NE OTC 
Date netted Marks marked Fresh 8 Jul 5 Jul 28 Jun 21Jun 14 Jun 
8July 119 28 °& 91 13 14 6 4 4 
12 July> 52 37 15 6 23 6° Ade 2° 2e 


12 Julys 104 74 30 12 
* Patch completely cleared. 
’ Twice normal sampling effort (about 1 hour). 
°12 jul values multiplied by 2. 
4Inc tides 2 individuals not captured on the 8th. 
© Also caught on the 8th. 


situation with the marked individuals first captured on the 5th and 
then recaptured on the 8th. Individuals marked earlier (e.g., 28, 21 
and 14 June) seem to be “resident’’; that is, captured on both the 8th 
and 12th. Of the 30 “new” individuals, 12 (about 50%) were born 
within the patch (wings were in perfect condition, no scale loss or still 
soft) and 18 had immigrated into the patch, as had two individuals 
marked on 28 June. These were not present on the 8th but were caught 
again on the 12th. 


Discussion 


The present study of adult D. plexippus was conducted in a region 
where the insect breeds continuously throughout the year. The host 
plants of D. plexippus, milkweeds, represent a discontinuous, hetero- 
geneous resource for adults and larvae. Milkweeds grow in dense, rel- 
atively well defined stands or patches. Patches vary greatly in area, 
plant density, age, condition and dispersion. The number of adults 
within any one patch varies through the year, the fluctuations repre- 
senting changing birth, death and movement rates. Adult monarchs 
are highly vagile and have the potential to range widely during the 
course of their lifetime. The evidence for monarch movement is ex- 
tensive and consists of: (1) the recapture of marked individuals at a 
patch other than the site of first capture—although the number of these 
in this study was small (due mainly perhaps to low sampling intensity), 
they indicate that monarchs can move between patches; (2) over half 
of the “new” individuals entering a patch population were not neces- 
sarily born there; (3) monarchs can be observed flying in all types of 
habitat and new milkweed patches are rapidly colonized (see Zalucki 
& Kitching, 1982a, b, c); and (4) various studies in North America in 
all seasons (e.g., Gibo & Pallett, 1979; Urquhart, 1960; Urquhart & 
Munger, 1970; Urquhart & Urquhart, 1976). 

The monarch’s vagility makes it difficult to study patch populations 


VOLUME 38, NUMBER 3 ea ler 


using techniques of mark-recapture. One difficulty is that in sampling 
a patch one is sampling a sub-population of a much larger population. 
Individuals in the sub-population move over a much wider area than 
is contained by a patch. For this reason marked individuals are not 
equally available for recapture in the sub- and parent populations. 
Movement into and out of the patch will cause fluctuations in the 
proportion of recaptures leading to large fluctuations in estimates of 
population size and large variances in associated vital rates (see also 
Brussard & Ehrlich, 1970). 

Movement into and out of patches seems to depend partly on the 
interactions between males and females as evidenced by changing sex 
ratios. The changing sex ratios could also be due to differential male- 
female mortality. The mark-recapture method does not distinguish 
between losses due to death and those due to emigration. However, 
from laboratory studies, male and female survivorship curves are iden- 
tical (fig. 6 in Zalucki, 1981b; x?,, = 0.908). This leaves differential 
movement rates and consequent patch residence times (see Table 8) as 
the most likely explanation for changing sex ratios. Certainly, male- 
female interactions are dramatic. Males pursue any flying object within 
range. Females will attempt to evade males, and high speed chases 
usually ensue (see Pliske, 1974, for detailed description of courtship 
behavior). We suggest these sexual interactions between males and 
females could provide a mechanism whereby females are differentially 
dispersed from a local population, such as exists within a patch. Shapiro 
(1970) provides evidence for a similar dispersal mechanism in pierids. 

The change in sex ratio towards females in early autumn (Fig. 8) is 
presumed to be due to the movement of females from around the 
countryside in towards large patches. It may even represent the return 
of individuals from inland areas, as the monarch’s summer range con- 
tracts (Smithers, 1977). (Eanes & Koehn, 1978 note a similar change 
in sex ratios for migratory populations in North America.) However, 
this would imply that the inland population has a skewed sex ratio, for 
which there is no evidence. 

Our observations indicate strongly that populations of monarchs 
around patches exhibit high rates of turnover. Our data on how far 
monarchs fly, given they leave a patch, are scant. Only five individuals 
were caught at sites other than the site of marking. These indicate the 
potential for cross-country movements but not what percentage of 
individuals undertake such movements. Of the butterflies netted, 87% 
are not recaptured (Table 1). Where do they go? Either monarchs 
“loiter” outside milkweed patches, moving in and out of local patches 
with some unknown percentage undertaking longer movements; or 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


most monarchs range widely if only by diffusion among many clumps 
of patches. Further intensive studies, some of which are in progress, 
will be needed to distinguish among these and other hypotheses. 


ACKNOWLEDGMENTS 


The authors with to thank all those who helped with the patch clearing exercise. M.P.Z. 
was supported by a CPRA scholarship and R.L.K., in part, by a grant from the A.R.G.C. 


LITERATURE CITED 


BRUSSARD, P. F. & P. R. EHRLICH. 1970. The population structure in Erebia epipsodea 
(Lepidoptera: Satyrinae). Ecology 51:119-129. 

DETHIER, V. G. 1959. Food plant distribution and density and larval dispersal as factors 
affecting insect populations. Can. Entomol. 91:581-596. 

EANES, W. F. & R. K. KOEHN. 1978. An analysis of genetic structure in the monarch 
butterfly Danaus plexippus L. Evolution 32:784-797. 

EHRLICH, P. R. 1965. The population biology of the butterfly, Euphydryas editha. I. 
The structure of the Jasper Ridge colony. Evolution 19:327-336. 

& L. E. GILBERT. 1973. Population structure and dynamics of the tropical 
butterfly Heliconius ethilla. Biotropica 5:69-82. 

Gipo, D. L. & M. J. PALLETT. 1979. Soaring flight of monarch butterflies, Danaus 
plexippus (Lepidoptera: Danaidae), during the late summer migration in southern 
Ontario. Canad. J. Zool. 57:1393-1401. 

JOLLY, G. M. 1965. Explicit estimates from capture-recapture data with both death 
and immigration—stochastic model. Biometrika 52:225-247. 

MUNGER, F. & T. T. HARRISS. 1970. Laboratory production of the monarch butterfly 
Danaus plexippus. J. Res. Lepid. 8:169-176. 

PLISKE, T. E. 1974. Courtship behaviour of the monarch butterfly, Danaus plexippus 
L. Ann. Entomol. Soc. Am. 68:143-151. 

SEBER, G. A. F. 1973. The Estimation of Animal Abundance. Griffin and Co. Ltd., 
London. 506 pp. 

SHAPIRO, A. M. 1970. The role of sexual behaviour in density related dispersal of pierid 
butterflies. Amer. Nat. 104:367-372. 

SHARP, M. A. & D. R. Parks. 1973. Habitat selection and population structure in 
Plebejus saepiolus Boisduval (Lycaenidae). J. Lepid. Soc. 27:17—22. 

SINGER, M. C. 1971. Evolution of food-plant preferences in the butterfly Euphydryas 
editha. Evolution 25:383-389. 

SMITHERS, C. N. 1973. A note on length of adult life of some Australian butterflies. 
Aust. Entomol. Mag. 1:62—66. 

1977. Seasonal distribution and breeding status of Danaus plexippus in Aus- 
tralia. J. Aust. Entomol. Soc. 16:175-184. 

SOUTHWOOD, T. R. E. 1966. Ecological Methods. Methuen, London. 

URQUHART, F. A. 1960. The Monarch Butterfly. Univ. Toronto Press. 

& F. MUNGER. 1970. A study of a continuously breeding population of Danaus 

plexippus. J. Res. Lepid. 7:169-181. 

& N. R. URQUHART. 1976. A study of the peninsular Florida populations of 
the Monarch butterfly (Danaus p. plexippus; Danaidae). J. Lepid. Soc. 30:73-87. 

ZALUCKI, M. P. 198la. Temporal and spatial variation of parasitism in Danaus plex- 
ippus (L.) (Lepidoptera: Nymphalidae: Danainae). Aust. Entomol. Mag. 8:3-8. 

1981b. The effects of age and weather on egglaying in Danaus plexippus L. 

(Lepidoptera: Danaidae). Res. Pop. Ecol. 23:318-327. 

198lc. Animal movement and its population consequences with a case study 

of Danaus plexippus L., Ph.D. Thesis, Griffith University, Australia. 

, A. CHANDICA & R. L. KITCHING. 1981. Quantifying the distribution and 

abundance of an animal’s resource using aerial photography. Oecologia (Berl.) 50: 

176-183. 


























VOLUME 38, NUMBER 3 219 





& R. L. KITCHING. 1982a. The dynamics of oviposition of Danaus plexippus 
(Insecta: Lepidoptera) on milkweed (Asclepias spp.). J. Zool. Lond. 198:103-116. 
1982b. Temporal and spatial variation of mortality in field populations of 
Danaus plexippus L. and D. chrysippus L. larvae (Lepidoptera: Nymphalidae). 
Oecologia (Berl.) 53:201-207. 


1982c. The analysis and description of movement in adult Danaus plexippus 
L. (Lepidoptera: Danainae). Behaviour 80:174-198. 





Journal of the Lepidopterists’ Society 
38(3), 1984, 220-234 


BUTTERFLIES OF TWO NORTHWEST 
NEW MEXICO MOUNTAINS 


RICHARD HOLLAND 
1625 Roma NE, Albuquerque, New Mexico 87106 


ABSTRACT. This article tabulates butterflies taken in the Chuska and Zuni Mts. of 
northwest New Mexico and extreme northeast Arizona. Emphasis is on the author’s own 
experience during the period 1971-1978. 


This paper is the third in a series which eventually will treat the 
butterfly fauna of all the major mountain ranges in New Mexico, except 
those which are unbroken extensions of the Colorado Rockies. The first, 
published by Holland (1974) dealt with six ranges in the central part 
of the state. The second, published by Ferris (1976) reported on the 
Grant-Catron County area, including the extensive Gila Mts. The pres- 
ent work is devoted to the Zuni and Chuska Mts. of northwestern New 
Mexico (see Figs. 1-3). Surveys of the butterflies of the Sacramento, 
Capitan, and Organ Mts. in southcentral New Mexico have also been 
completed, but publication of this work is being delayed until the 
Guadalupe Ridge has been investigated as well. Additionally, a survey 
of the Jemez Mts. in northcentral New Mexico is now completed and 
will be published soon. Besides the Guadalupe Ridge, studies of several 
ranges outlying the Gila are planned. These ranges include the Black, 
Animas and Datil Mts. 

About four quite limited regional lists of the New Mexico fauna 
have also appeared; these lists are catalogued in the above-mentioned 
Holland and Ferris articles. Additionally, there is an unpublished dis- 
sertation by Toliver (1978) which tabulates every butterfly specimen 
taken in New Mexico prior to 1978. 

The Chuska Mts. fall across the Arizona—New Mexico state line, but 
this survey considers equally the fauna on both sides of the line; natural 
rather than political boundaries delineate the study areas. The Carrizo 
Mts. (see Fig. 1), however, are not included in the present study, as 
they lie entirely in Arizona and are isolated from the main part of the 
Chuskas by a considerable expanse of very arid low desert. 

The present article reports my studies for the years 1971-1978, dur- 
ing which IJ systematically surveyed the Zuni and Chuska Mts. While 
a third range, Mt. Taylor, also lies in this area (see Fig. 1), its fauna 
has already been reported by Holland (1974) and will not be redocu- 
mented here. 

Endemism of butterfly species is rare in the New Mexico mountains. 
In this respect, the northwest part is typical of the state as a whole. 
Mt. Taylor and the Zuni Mts. support no endemics at any taxonomic 


VOLUME 38, NUMBER 3 221 








| COLORADO 


CARRIZO 


NEW MEXICO 






SAN JUAN CO. RIO ARRIBA CO. 






CHUSKA 
MOUNTAINS 











Mc KINLEY CO. SANDOVAL CO. 


APACHE CO. 


VALENCIA CO. 


20 Mi 








Fic. 1. Map of the mountains of northwestern New Mexico, showing the 8000’ ele- 
vation contours. 


level. The Chuskas have a single subspecific endemic, Occidryas anicia 
chuskae Ferris & R. Holland. This near-total absence of endemism 
indicates that the desertification of the New Mexico lowlands is ex- 
tremely recent, geologically speaking, in comparison with, for instance, 
the Mojave. I believe New Mexico mountains have been isolated for 
less than 4000 years. The one endemic taxon seems more likely to be 
a relict than something which actually evolved on the Chuskas. 


222 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The format of my earlier article was different from that of Ferris; 
Ferris gave more precise and detailed data which permitted cross- 
referencing to counties as well as mountain ranges. In this article, 
Ferris’ format will be used. 


Localities 


The Chuska Mts. are sedimentary (red sandstone and limestone). 
The Chuska Mts. mainly consist of a vast plateau with sharp escarp- 
ments on all sides. Scenically, the red sandstone cliffs and canyons are 
very dramatic. The plateau has numerous subsidences where natural 
lakes have formed, some of which cover hundreds of acres. Much of 
the plateau top is forested with aspen (Populus tremuloides Michx.) 
and Douglas fir (Pseudotsuga taxifolia Mayr). There are also large 
open meadows. The southern end of the Chuska plateau is somewhat 
drier and lower, and ponderosa pine (Pinus ponderosa Laws.) is dom- 
inant. Collecting is generally not good on the plateau itself; the canyons 
running off the plateau tend to have a much richer fauna. Flora which 
significantly affects the butterfly diversity includes several willows (Sa- 
lix spp.), Gamble oak (Quercus gambelii Nutt.), several species of ju- 
niper, several Yucca spp., Ceanothis fendleri Gray, Rumex sp. (pos- 
sibly introduced), several columbines (Aquilegea spp.), several 
Penstemon spp. and paintbrushes (Castilleja sp.), cliffrose (Cowania 
mexicana D. Don.), and at least five species of Eriogonum. Lower 
elevations have considerable stands of saltbush (Atriplex sp.) and sage 
(Artemisia sp.). The fauna is undoubtedly affected by the absence of 
some plants as well. Missing flora include locust (Robinia sp.), hack- 
berry (Celtis sp.), walnut (Juglans sp.), Agave sp., and mesquite (Pro- 
sopis sp.). 

The Chuska Mts. lie entirely on the Navajo Indian Reservation. At 
present, the tribal authorities are not hostile to outsiders, and nearly 
the entire reservation is open to the general public without written or 
oral permission. This pleasant situation will probably change. On the 
negative side, sheep and goats have been allowed to devastate most of 
the Chuskas. It is unlikely that any Lepidoptera have actually been 
exterminated, but many species tend to be scarce and local because of 
the land abuse. Also, poorly regulated lumbering has been permitted 
over wide areas without even the minimal erosion-control and under- 
story protection efforts one usually sees in national forests. Additionally, 
roads are terrible, and getting into the canyons where collecting is good 
tends to be very challenging. Regrettably, there is an element of sus- 
picion that false Lepidoptera records have been claimed for the Chus- 
kas, and to a lesser extent, the Zunis. 

The Zuni Mts. are more rolling with few dramatic canyons or es- 


VOLUME 38, NUMBER 3 223 


carpments. In contrast to the Chuskas, the Zunis are principally vol- 
canic. The flora is surprisingly similar to the Chuskas, considering the 
geological differences. The greatest disparity is that oak and ponderosa 
pine are more prevalent, with aspen and Douglas fir correspondingly 
restricted. Good collecting in the Zunis is less limited to the lower 
canyons. The Zuni Mts. have also been subjected to excessive land use, 
although most of the grazing is cattle rather than sheep and goats, so 
the destruction is less radical than in the Chuska Mts. The Zuni Mts. 
are mostly in the Cibola National Forest. Consequently, more conser- 
vative lumbering techniques have been practiced than in the Chuska 
Mts. Also, secondary roads are better maintained in the Zunis. Although 
the Zunis have been abused less than the Chuskas, it appears Speyeria 
nokomis (W. H. Edwards) has suffered extinction in the Zunis. 

This land is subject to extreme temperatures, especially in winter. 
Summer temperatures reach 38°C; winter lows colder than —50°C 
have been recorded at the reservation town of Zuni. I have no idea 
how cold it gets in Roof Butte and Mt. Sedgwick. Precipitation occurs 
mainly in December—March and July-September. May is the driest 
month. Annual precipitation varies from 0.2 m in the rain shadow on 
the New Mexico side of the Chuskas, to around 0.8 m on Roof Butte. 

Specific collecting sites and their alphanumeric codes are given be- 
low. The locality code symbols appear on maps in Figs. 2 and 8. As 
stated previously, this style of data presentation is copied from Ferris 
(1976). 


ZUNI MTS., McKINLEY COUNTY, NEW MEXICO. Clo-Chen-Toh Ranch (CCT) 
7100’; Cottonwood Gulch (CG) 7500’; Ft. Wingate (FW) 6800’; Gallup (G) 6700’; Grass- 
hopper Canyon (GC) 7500’—7700'; Grasshopper Spring (GS) 7500’; McGaffey (M) 7500’; 
McGaffey Lake (ML) 7500’; Milk Ranch Canyon (MRC) 7600’; Nutria Diversion Reser- 
voir (NDR) 7300’; NM Rt. 58 at Jct. to NM Rt. 32 (NMJ) 6600’; NM Rt. 58 at Jct. to 
Nutria (NJ) 7000’; Prewitt Tank (PT) 7800’; Ramah (R) 7000’; Ramah Lake (RL) 7000’; 
Stinking Spring (SS) 7500’; Wingate Tank (WT) 7600’; NM Rt. 400 at Jct. to Interstate 
40 (400) 6900’; NM Rt. 412 at Jct. to Interstate 40 (412) 7200’. 

ZUNI MTS., VALENCIA COUNTY, NEW MEXICO (now in CIBOLA COUNTY by 
action of the New Mexico legislature after this study was completed). Lower Bluewater 
Canyon (IBC) 7000’-7300' (below Bluewater Dam); upper Bluewater Canyon (uBC) 
7300'-7500' (above Bluewater Dam); Bluewater Dam (BD) 7300’; Bluewater Village (BV) 
6500’; Cebolla Canyon (CC) 7000’; Diener Canyon (DC) 8000'—9000’; El Morro National 
Monument (EMNM) 7000’; Kettner Canyon (KC) 8000’; Log Cabin Canyon (LCC) 7500'- 
8000’; Manga Canyon (MC) 7000’; Mt. Sedgwick (MS) 9300’; Ojo Redondo (OjR) 8000’; 
Oso Ridge (OsR) 8900’; Pink Rose Canyon (PRC) 7000’; Pole Canyon (PoC) 8000’—-8500’; 
Prop Canyon (PrC€) 8000’; Sawyer (S) 8000’; San Rafael (SR) 6700’; Tusas Mesa (TM) 
8000’; lower Zuni Canyon (IZC) 7000’-7500'; upper Zuni Canyon (uZC) 8000'-8200’. 

CHUSKA MTS., McKINLEY COUNTY, NEW MEXICO. Chuska Peak (CP) 8700’; 
Navajo (N) 6800’; Tohatchi (Toh) 6000’—7500’; Tohatchi Lookout (TL) 8300’; Whiskey 
Lake (WsL) 8000’. 

CHUSKA MTS., SAN JUAN COUNTY, NEW MEXICO. Big Gap Reservoir (BGR) 
5000’; Beautiful Mountain (BM) 8000’; Cox Canyon (CC) 6200’; Occidryas anicia chuskae 
Type Locality (OacTL) 7700’; FAA Installation (FAA) 9000’; Owl Springs (OS) 7500’; 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


400 





McKINLEY Co 
NM 


GRANTS 


NJ 


ZUNI 


VALENCIA Co 
(CIBOLA Co) 


NM 10 MI 





Fic. 2. Detail of the Chuska Mts., showing the 8000’ elevation contours and collecting 
localities. 


Sanostee (Sn) 5500’; Shiprock Mt. (SM) 5500’; Toadlena (Toa) 6500’; Wheatfields Creek 
(WfC) 7400'-8200'; Whiskey Creek (WhC) 7700’; Washington Pass (WP) 8000’. 

CHUSKA MTS., APACHE COUNTY, ARIZONA. Buffalo Gap (BG) 7800’; Chinle 
(Ch) 5500’; Cove (Co) 7000’-8000'; Ganado (G) 6500’; Hunters Point (HP) 7000’; Luka- 
chukai (L) 7000’; Luka Peak (LP) 9200’; Roof Butte (RB) 8500'—9600’; Red Lake (RL) 
6000’; Red Rock (ReR) 6000'—7000’; Round Rock (ReR) 5500’; Sawmill (Sw) 7800’; Spider 
Rock (SR) 7500’; Tsale Creek (TC) 7500’-8500'; Wheatfields Lake (WfL) 7000’; Wagon 
Wheel Campground (WWC) (Lukachukai Creek) 7500’; AZ Rt. 264 at Jct. to Sawmill, 
AZ (264) 7800’; 8 mi. north of Wheatfields Lake on Tsale Cr. (GBMNWE£L) 7500’. 


Checklist 


In the following checklist, mountain range and localities are noted 
as well as the flight period. (A “+” before a date indicates a common 
species which flies considerably later than the date indicated but for 
which late-season specimens were merely observed, not collected.) No- 
menclature and species number is that of Miller and Brown (1981) 
except in a few cases where my opinion is strongly different. 

Collectors, besides myself, who have supplied records are Richard 
Bailowitz (RB), Robert Langston (RL), James Scott (JS), Michael Fisher 


VOLUME 38, NUMBER 3 225 








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Fic. 3. Detail of the Zuni Mts., showing the 8000’ elevation contours and collecting 
localities. 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(MF), Clifford D. Ferris (CDF), Michael Toliver (MT), Kilian Roever 
(KR), John Justice (JJ), Carl Cushing (CC), Oakley Shields (OS), Bruce 
Griffin (BG), and Ray E. Stanford (RES). Records from the Toliver 
manuscript are denoted (TM). Most of these records are very old and 
due to John Woodgate’s collecting at Ft. Wingate; hence, if known, 
the year of capture is included with (TM) records. Where a date is 
replaced by a “P” in (TM) records, the record probably is of the period 
1906-1911. The Woodgate material from Ft. Wingate is now part of 
the American Museum of Natural History collection except for ly- 
caenids which apparently are mostly in the Carnegie Museum. 

John Woodgate was a fence-rider who carried a butterfly net until 
his eyesight failed. We have records of his at Ft. Wingate, NM, from 
1906 until 1911, and at Jemez Springs, NM, from 1912-19138 (Williams, 
1914); after this he apparently vanished. From the diversity of his 
records, it is obvious that he had become a much more sophisticated 
collector by 1913 than when he started in 1906. As recently as 1970, 
some of his material was under glass in a bar in Jemez Springs. 

Conditions at Ft. Wingate are very different than they were in 
Woodgate’s day, and some species have probably disappeared from 
the immediate vicinity. In fact, the town itself has been moved nine 
miles east! The pre-1912 Ft. Wingate records presumably refer to the 
old site. 


HESPERIIDAE-PYRGINAE 


7. Epargyreus clarus (Cramer). Chuska Mts.: ? no specific data (KR). To my knowl- 

edge, the foodplant Robinia is absent from Chuska and Zuni Mts. 

20. Zestusa dorus (W. H. Edwards). Chuska Mts.: ? no specific data (KR). 

48. Thorybes pylades (Scudder). Found almost everywhere in study area above 7000’. 
May 4-July 8. 

50c. Thorybes mexicana dobra Evans. Chuska Mts.: WP, FAA, vic. WfL, 10 mi. E 
WEL on WIC. May 19-July 8. 

83. Erynnis icelus (Scudder & Burgess). Chuska Mts.: WWC, RB, 8 mi. N WfL on 
TC, 12 mi. N WfL on TC, 10 mi. E WfL on WEC. June 19-July 8. 

84a. Erynnis brizo burgessi (Skinner). Chuska Mts.: RB, 2 mi. SW Cove, 2 mi. SE Cove, 
8 mi. N WfL on TC, 12 mi. N WfL on TC, BG, WWC, 2 mi. NW Toh, CP, 
WP, BM. Zuni Mts.: NDR, SS, MS, IZC, FW (TM, 1908 & 1909). April 23- 
July 4. 

86. Erynnis telemachus Burns. Chuska Mts.: 2 mi. NW Toh, Toa, WWC, WP, 4 mi. 
W WP, BG, 12 mi. S Toa, 12 mi. N WfL on TC, RB, 2 mi. SE Co, 10 mi. S SR. 
Zuni Mts.: 1BC, 4 mi. S FW, GS, IZC, RL, NMJ, FW (TM, 1908 & 1909). April 
19-+June 11. 

90. Erynnis horatius (Scudder & Burgess). Chuska Mts.: ? no specific data (KR). 

93a. Erynnis pacuvius pacuvius (Lintner). Chuska Mts.: RB, 4 mi. S Co, TL, 2 mi. NW 
Toh, TL. Zuni Mts.: MS, TM, GS, PoC, DC, IZC, PrC, FW (TM, 1908 & 1909), 
OsR (CC). May 2-July 4. 

95. Erynnis funeralis (Scudder & Burgess). Chuska Mts.: Toa, WfL. Zuni Mts.: WT 
(KR & RH). May. 

98. Erynnis afranius (Lintner) & persius (Scudder). Chuska Mts.: BM, WhC, WWC, 
RB, WP, RL, BG, LP, 6 mi. SW Sn, 2 mi. SE Co, 2 mi. SW Co, 12 mi. S Toa, 


VOLUME 38, NUMBER 3 DO 


102. 


103. 


104. 


107. 


109. 


115. 


118a. 


121. 


145. 


146. 


152. 


155. 


156a. 


157. 


158n. 


159. 


168a. 


166. 


175a. 


12 mi. N WEL on TC, TL. Zuni Mts.: PoC, MS, PrC, uBC, DC, KC, uZC, GC, 
IZC, GS, PRC, OsR, FW (TM, 1909). April 24—August 18. 

Pyrgus xanthus W. H. Edwards. Chuska Mts.: ? no specific data (KR). Zuni Mts.: 
2 mi. W M, DC, 8 mi. N OjR, OjR, BD, PoC, PrC, IZC. April 25-May 30. 
Univoltine. 

Pyrgus scriptura (Boisduval). Chuska Mts.: ReR. Zuni Mts.: PoC, SS, NJ. May 6- 
August 19. At least bivoltine. 

Pyrgus communis (Grote) & albescens Plotz. Most material from the study area 
is probably referable to communis. However, a somewhat mosaic distribution 
exists, with certain pockets of albescens being existent, especially around BD 
(Toliver, 1978). I am not convinced two species are involved here. I do not think 
the concept of a single genetically dimorphic species should be ruled out in cases 
such as this and Celotes in the absence of corroborating fertility studies. In 
general, specimens are found nearly everywhere in the study area except above 
8000’. At least April-September. 

Pyrgus philetas W. H. Edwards. Chuska Mts.: L (CDF & RH). June 26. This is 
definitely not a resident species. 

Heliopetes ericetorum (Boisduval). Chuska Mts.: HP, Toa. Zuni Mts.: NJ (KR & 
RH), FW (TM, 1910). June 10-July 3. Probably migratory and not present every 
year. 

Pholisora catullus (Fabricius). Zuni Mts.: ZC, IBC, FW (TM, 1906). June 27-July 
30. 

Pholisora alpheus alpheus (W. H. Edwards). Chuska Mts.: 2 mi. E WP, BGR. Zuni 
Mits.: NDR, FW, NJ, 400, 6 mi. E R. May 5-July 3. 

Piruna pirus (W. H. Edwards). Chuska Mts.: WWC, RB, BG, W of Sn at 8000’, 
8 mi. W Toa, 10 mi. E WfL on WIC. June 22-July 21. 


HESPERIIDAE-HESPERIINAE 


Oarisma garita (Reakirt). Zuni Mts.: IZC, uBC, MS, ML, LCC, PoC. June 24- 
August 6. 

Oarisma edwardsii (Barnes). Zuni Mts.: NDR, PoC, GS, LCC, MRC, PRC, NJ, 
FW (TM, 1908), EMNM (RL). July 2-August 6. 

Yoretta rhesus (W. H. Edwards). Zuni Mts.: PoC, BD, 4 mi. E R, NDR, 3 mi. W 
M, LCC, PT, FW (TM, 1907 & 1908). May 21-June 15. Univoltine. 

Stinga morrisoni (W. H. Edwards). Chuska Mts.: RB, TL, CP, 3 mi. S WhL, WP, 
264. Zuni Mts.: PoC, MS, uZC, IZC. May 1-June 18. 

Hesperia uncas uncas W. H. Edwards. Chuska Mts.: WfL, Toa, 3 mi. W Toa, 2 
mi. NW Toh, BM, HP. Zuni Mts.: NDR, NMJ, PoC, IZC, uBC, 412, 400, NJ, 
FW, GC, 12 mi. S G, PT. June 8—August 22. Bivoltine. 

Hesperia juba (Scudder). Chuska Mts.: L (JJ & RH). May 18. 

Hesperia comma susanae L. Miller. Chuska Mts.: ? WP (KR). Zuni Mts.: SS, 4 mi. 
SW FW. August 26-September 5. 

Hesperia woodgatei (R. C. Williams). Chuska Mts.: 2-4 mi. NW Toh. Zuni Mts.: 
2 mi. S FW. September 10—+September 24. Woodgate also recorded this species 
at FW in 1907 and 1908, several years before the type series was taken (1913) 
at Jemez Springs, NM. 

Hesperia pahaska pahaska (Leussler). Chuska Mts.: 2 mi. NW Toh, RL, TL, CP, 
3 mi. S WhL, WP, BM, HP. Zuni Mts.: 2 mi. S FW, GS, PoC, SS, DC, WT, IBC, 
LCC, PT. May 17-+July 4. 

Hesperia viridis (W. H. Edwards). Chuska Mts.: Toa, BG, 2 mi. E CP. Zuni Mts.: 
LCC, MG, IZC, PRC, PoC. May 29-July 28. 

Polites sabuleti sabuleti (Boisduval). Chuska Mts.: Toa. May 26. A curious situation 
exists here from two viewpoints. Inspection of a particular arroyo perhaps 15 
times over eight years yielded nothing. However, on 26 May 1978, sabuleti was 
present in numbers. Subspecies chusca (W. H. Edwards) apparently was not 
described from the Chuska Mts., but from Mohave Co., Arizona (Brown & 


228 


Tle 
igh 


187a. 
194. 
199. 


2a 


222. 
225. 
230. 


231. 


233. 
236. 


249. 


286c. 


289. 


297a. 
3038a. 


308. 


3lla. 


312b. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Miller, 1980). “Chuska” is a Navajo word meaning “white fir.” “White fir” 
(Abies concolor (Gordon & Glendinning)) occurs in many Arizona mountain 
ranges including the Hualapais of Mohave Co., but not in the Chuskas (Kearney 
& Peebles, 1964; Pearce, 1975). 

Polites draco (W. H. Edwards). Chuska Mts.: ? no specific data (KR). 

Polites themistocles (Latreille). Chuska Mts.: Wf£L, WWC, Toa, 8 mi. N WfL on 
TC, OacTL, WhC, 4 mi. E WfL, 2 mi. NW Toh. Zuni Mts.: KC, PT. June 8- 
August 5. 

Atalopedes campestris campestris (Boisduval). Chuska Mts.: WhC. August 5. 

Ochlodes snowi (W. H. Edwards). Chuska Mts.: ? no specific data (KR). Zuni Mts.: 
PrC, 4 mi. S FW, GS, IZC, PoC. July 29—August 15. 

Poanes taxiles (W. H. Edwards). Chuska Mts.: Toa, 2 mi. E CP. Zuni Mts.: 4 mi. 
N NMJ, GS, PoC, LCC, MC, IZC, NDR, FW (TM, ?). June 8-July 28. 

Euphyes ruricola ruricola (Boisduval). Chuska Mts.: 1 mi. W Toa, WWC, 8 mi. 
N WEIL on TC, Toa, 12 mi. N WfL on TC, 8 mi. E WfL on WEC. June 8- 
August 5. 

Atrytonopsis vierecki (Skinner). Chuska Mts.: Toa, Sn. Zuni Mts.: IZC, NJ, LCC, 
SR, MC, FW (TM, 1907). May 19-June 8. 

Atrytonopsis python (W. H. Edwards). Zuni Mts.: LCC, MC, PoC, NJ, SR, MRC, 
BD (MT), FW (TM, 1907). May 29-July 2. 

Amblyscirtes cassus W. H. Edwards. Zuni Mts.: POC, LCC, PRC, GS, IZC, FW 
(TM, ?). June 25-July 28. 

Amblyscirtes aenus aenus W. H. Edwards. Chuska Mts.: ? no specific data (KR). 
Zuni Mts.: IZC, GS, LCC, NDR, PRC, GC, PoC, FW (TM, 1907). June 1—August 
8. An undescribed subspecies of aenus with very whitish dorsal forewing spots 
occurs in southern New Mexico. The present populations are typical. 

Amblyscirtes oslari (Skinner). Zuni Mts.: GS. June 19. 

Amblyscirtes texanae Bell. Zuni Mts.: OsR. August 18. One very worn specimen 
taken. Undoubtedly a stray from at least 100 miles to the south or west. 

Amblyscirtes phylace (W. H. Edwards). Zuni Mts.: POC, GS, IZC, PoC, DC, BC, 
MRC. May 30-July 29. 


MEGATHYMIDAE 


Megathymus coloradensis navajo Skinner. Chuska Mts.: 2 mi. N BG, 2 mi. NW 
Toh, WWC, 2 mi. SE Co, L, 12 mi. S Toa. Zuni Mts.: PoC, GS, 3 mi. NE M, 
IZC, FW. April 29-June 9. Woodgate took the type series of navajo at FW in 
1910 or 1911 (Skinner, 1911). 

Megathymus streckeri (Skinner). Chuska Mts.: 6 mi. SW of Jct. of Nav. Rt. 34B 
& main rd. to Sn, BM. Zuni Mts.: FW (TM, ?). May 20-July 4. Curiously, all 
wild-caught specimens of coloradensis have been males and all streckeri have 
been females. 


PAPILIONIDAE 


Battus philenor philenor (Linnaeus). Chuska Mts.: 2 mi. SE of Co (visual record 
by RH and CDF). 25 June 1978. 

Papilio polyxenes asterius Stoll. Chuska Mts.: 3 mi. NW LP, TL, Toa. Zuni Mts.: 
PT, NJ, BD (OS). May 21-July 23. 

Papilio bairdii W. H. Edwards. Chuska Mts.: 8 mi. SW Sn, BG, CP, LP. Zuni Mts.: 
NDR, 4 mi. S FW, GS, MS, WT, PoC, ML, BD (TM, 1969, leg. Funk). May 2- 
September 14. 

Papilio zelicaon zelicaon Lucas. Chuska Mts.: TL, FAA, WP, RB, WWC, WIL, 4 
mi. S Co. Zuni Mts.: MS, GS, PoC, NDR, DC. April 29-June 14. Univoltine 
population. 

Papilio indra minori Cross. Zuni Mts.: FW (collector unknown, specimen is in 
LACM). 27 May 1917. 


VOLUME 38, NUMBER 3 229 


23la. Pterourus rutulus rutulus (Lucas). Chuska Mts.: WWC. Zuni Mts.: PoC, NDR. 
May 21-June 14. Unaccountably rare in Chuskas and Zunis. 

322. Pterourus multicaudata (W. F. Kirby). Chuska Mts.: RL, 3 mi. NW Co. Zuni Mts.: 
15 mi. S Grants on NM 53, PoC, EMNM (RL, sight record). June 24-July 23. 
This species is also unaccountably rare in the Chuskas and Zunis. 


PIERIDAE 


329. Neophasia menapia (C. & R. Felder). Chuska Mts.: 2 mi. W Toa, 1 mi. E WP, 8 
mi. N WfL on TC, Toa, OacTL. Zuni Mts.: GS, PoC. July 16-August 7. 

332. Pontia beckerii (W. H. Edwards). Chuska Mts.: BGR, 2 mi. S SM, L, 6 mi. S Sn, 
Toa, SM, 2 mi. E RoR. May 5-July 24. At least bivoltine. 

338c. Pontia sisymbrii elivata (Barnes & Benjamin). Chuska Mts.: 2 mi. E of Toh, 2 mi. 
SE Co, WWC, WP, 10 mi. SW Toa, FAA, RB, 7 mi. SW Toa, WfL, TL, Toa, 
SR. Zuni Mts.: MS, BD, IZC, NDR, OsR (CC). April 22-June 14. 

334. Pontia protodice (Boisduval & LeConte). Found everywhere, especially below 
7500’, in the study area. Some phenotypes from the study area fall into the 
outdated concept (Brown, 1957) of occidentalis (Reakirt), but not the present 
concept (Chang, 1963). At least from April to September. 

338. Artogeia rapae (Linnaeus). Chuska Mts.: Ch, Toa (MF & RES), LP. Zuni Mts.: 
BV, SR. May 11-+June 26. 

348c. Euchloe hyantis lotta Beutenmuller. Chuska Mts.: FAA, 7 mi. SW Toa, 2 mi. SW 
Co, TL. April 23-June 4. 

348b. Anthocharis sara inghami Gunder. Chuska Mts.: 2 mi. SE Co, Toa, 7 mi. SW Toa. 
Chuska populations grade towards julia W. H. Edwards. Zuni Mts.: MS, uBC, 
4 mi. S FW, GC, PrC, PoC, 3 mi. NE M, IZC, NDR. April 21—May 14. 

35l1c. Colias philodice eriphyle W. H. Edwards. Chuska Mts.: WfL, Toa, 5 mi. S RB, 
BG, 6 mi. W Sn, 8 mi. N WfL on TC, 16 mi. N WfL on TC. Zuni Mts.: BV. 
May 15-September 2. 

352. Colias eurytheme Boisduval. Found almost everywhere in NM. Records from 
Zunis and Chuskas are restricted to April-September due to collecting season. 
Sandia Mts. (Albuquerque, NM) records exist for every month. 

355. Colias alexandra W. H. Edwards. Chuska Mts.: ? BG (KR). 

368a. Zerene cesonia (Stoll). Chuska Mts.: RL, L, 8 mi. N WFL on TC. Zuni Mts.: GS. 
May 12-July 30. 

371b. Phoebis sennae eubule (Linnaeus). Zuni Mts.: R, GS, NMJ. August 6—-August 19. 
This species had strong northward migrations in 1976 and 1977. It is not a 
permanent resident of the study area. 

380. Eurema mexicana (Boisduval). Chuska Mts.: several sight records; no actual spec- 
imens of this migratory species. Zuni Mts.: WT, POC. At least May-July. 

388. Eurema nicippe (Cramer). Chuska Mts.: 16 mi. N WfL on TC, 8 mi. N WfL on 
TC, 10 mi. S SR. Zuni Mts.: WT, POC, RL, BD (MT). May 11 to at least July. 

389. Nathalis iole Boisduval. Chuska Mts.: Sn, Toa, WfL. Zuni Mts.: PoC, NMJ, GS, 
BD (MT). April 22—late summer. 


LYCAENIDAE 


392d. Tharsalea arota schellbachi Tilden. Chuska Mts.: ? no specific data (KR). Zuni 
Mts.: FW (TM, ?. Specimen is in Carnegie Museum. It may not be a Woodgate 
specimen. ). No date. 

404. Epidemia helloides (Boisduval). Chuska Mts.: WfL, 3 mi. W Toa. May 17-June 
22. Number of broods uncertain, but probably more than one. WfL (just below 
dam) is the site of the only known helloides population in Arizona. The Rumex 
at this site may be introduced. 

408a. Hypaurotis crysalus crysalus (W. H. Edwards). Chuska Mts.: 2 mi. SE Co, 2 mi. 
NW Co, Toa. Zuni Mts.: GS, FW (TM, 1907). July 3-August 13. Possibly bivol- 


230 


412a. 
418b. 
422c. 


433. 


446a. 


449b. 


452. 


458a. 


463. 


464d. 
465b. 


47a. 


478c. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


tine in some areas. Curiously, bivoltinism is suspected in more boreal parts of 
northcentral NM, and in the Sierra Madre of Chihuahua. Chuska and Zuni 
populations appear to be univoltine. 

Atlides halesus halesus (Cramer). Chuska Mts.: Toh, 2 mi. SE Co, Toa, 6 mi. SW 
S, CP. Zuni Mts.: GS, POC, FW (TM, 1906 & 1907). May 2—August 5. 

Satyrium behrii crossi (Field). Chuska Mts.: BG, 3 mi. SE Co, 3 mi. NW Co, 10 
mi. SW Sn, LP, 2 mi. SW Co. Zuni Mts.: GC, FW (TM, ?). June 25-July 18. 

Satyrium sylvinus itys (W. H. Edwards). Zuni Mts.: FW (TM, ?. Specimen is in 
Carnegie Museum. It may not be a Woodgate specimen.). No date. 

Ministrymon leda (W. H. Edwards). Chuska Mts.: L. Zuni Mts.: 5 mi. W M, LCC. 
May 21-June 19. These specimens are obviously migrants as the foodplant Pro- 
sopis does not grow in or near the Zunis or Chuskas. 

Callophrys apama apama (W. H. Edwards). Chuska Mts.: TL, WWC, 8 mi. W£L 
on WIC, 2 mi. NW of Toh. Zuni Mts.: PrC, GS, PoC, FW (TM, 1909 & 1910), 
OjR (RB). April 29-August 15. Bivoltine, peaks in May and early July. 

Callophrys sheridanii neoperplexa Barnes & Benjamin. Chuska Mts.: no actual 
records, but RES believes it may be present as it is found farther south in Apache 
Co., AZ (White Mts.). sheridanii has erroneously been reported as occurring in 
northeastern NM. The only known NM population is in the Sacramento Mts. in 
the southern part of the state, although a single specimen turned up in 1983 in 
the San Juan Mts. just south of the Colorado state line (JS & RES). Thus, its 
distribution is strongly disjunct. 

Mitoura spinetorum (Hewiston). Chuska Mts.: 2 mi. SE Co, RB, WP, TL, FAA, 
2 mi. NW Toh. Zuni Mts.: SS, PoC, DC, FW, IZC, NDR, GS, BD (MT). April 
23-August 14. Out-of-state collectors often express surprise that spinetorum is 
a common butterfly in New Mexico, as it apparently is much less abundant in 
other parts of its range. There is a record of 400 specimens being taken in a day 
near Magdalena, NM (JJ). 

Mitoura siva siva (W. H. Edwards). Chuska Mts.: WWC, Toa, 2 mi. NW Co, Co, 
L, Toh, 6 mi. SW Sn, 2 mi. E CP, 4.5 mi. S HP, 11 mi. SE RoR, LP. Zuni Mts.: 
PoC, CC, IBC, IZC, 4 mi. S FW,! NDR, RL, GS, SR, 6 mi. E R. April 22- 
August 6. 

Sandia mcfarlandi P. Ehrlich & Clench. Zuni Mts.: 1ZC. 2 May 1976 (2 specimens). 
Not seen in other years. In 1974, Holland stated that the foodplant Nolina texana 
does not occur on Mt. Taylor. Since then, I discovered a colony visible only by 
railroad about 10 miles east of Grants. Intensive searching has failed to turn up 
mcfarlandi at this site. The IZC population is extremely small, and may be on 
the verge of natural extinction (human impact on the area is minimal). A similar 
very weak population occurs near Acoma, about 30 miles east of Grants. No 
adults have ever been taken at Acoma, but a single larva was found there in 
1971 and reared out. 

Incisalia augustus iroides (Boisduval). Chuska Mts.: taken at CC (MF & RES), 29 
April 1974; no records from actual slopes of Chuskas. 

Incisalia fotis fotis (Strecker). Chuska Mts.: 1 mi. NW Toh, Toa, L. Zuni Mts.: 
PrC, PoC, GS, FW (TM, ?). April 21-June 8. 

Incisalia eryphon eryphon (Boisduval). Chuska Mts.: WWC, WP, 2 mi. NW Toh, 
BG, FAA, RB, 4 mi. W WP, 7 mi. SW Toa, WEL, 8 mi. N WfL on TC, 10 mi. 
S SR. Zuni Mts.: PrC, PoC, DC, 3 mi. W M, GS. April 24-June 14, 

Strymon melinus franki Field Chuska Mts.: 3 mi. W Sn, 8 mi. N WfL on TC, 2 
mi. SE Co, 11 mi. E RoR, 4 mi. N OS, WWC, BGR, L, 8 mi. SW Sn, 2 mi. NW 


‘Miller and Brown indicate the type locality of siva is “Probably near Fort Wingate, Arizona [sic].” Actually, Fort 
Wingate is in New Mexico, and always has been as Arizona was once part of New Mexico. For a general discussion of 
the indiscriminate use of the locality designation “Arizona” on material collected by the Wheeler Expedition, see Brown 
(1983). Refer to Brown and Opler (1970) for a discussion of the confusion caused by this indiscrimination with respect 
to fixing the siva type locality. 


VOLUME 38, NUMBER 3 231 


495. 


498. 


502a. 
504d. 


505h. 


508a. 


508b. 
5lle. 
511d. 


513d. 


514f. 


517b. 


518b. 


520b. 


522b. 


526e. 


544a. 


5950. 


Toh, 4.5 mi. S HP, 264, BM. Zuni Mts.: GS, ML, IZC, 5 mi. W M, NDR, MRC, 
6 mi. E R, FW (TM, 1907). May 11-+August 7. 

Brephidium exilis (Boisduval). Chuska Mts.: 264, RoR. Zuni Mts.: PoC, FW, 400, 
NMJ, G, RL. May 11-August 31. 

Leptotes marina (Reakirt). Chuska Mts.: WWC, Toa, 5 mi. E G, Ch, 10 mi. S SR, 
264. Zuni Mts.: NDR, 5 mi. W M, PoC, 1 mi. N FW, PT. May 11-—August 6. 
Hemiargus isola alce (W. H. Edwards). Common and universally distributed in 
NM at elevations up to 7500’. At least May—September in Zunis and Chuskas. 
Everes amyntula herrii F. Grinnell. Chuska Mts.: 2 mi. SE Co, Toa, WWC, 4 mi. 

S Co. Zuni Mts.: uZC, PoC, PrC, GS, IZC. May 1-August 15. 

Celastrina argiolus cinerea (W. H. Edwards). Chuska Mts.: BG, WWC, WP, FAA, 
4 mi. W WP, HP, WhC, 2 mi. SE Co, Toa, 4 mi. S Co, 2 mi. NW Toh. Zuni 
Mts.: PoC, 5 mi. W M, DC, PoC, NDR, FW (TM, 1906 & 1910). Form “mar- 
ginata’” occurs commonly in the Chuskas but never in the Zunis. April 30- 
August 9. 

Euphilotes battoides centralis (Barnes & McDunnough). Zuni Mts.: PoC, PrC, 
IZC, GC, 400, 5 mi. W M, 3 mi. NE M, OsR, PrC, MC, EMNM (RL). July 10- 
August 22. 

Euphilotes battoides ellisii (Shields). Chuska Mts.: 2-3 mi. NW RoR, 20 mi. S Ch 
on AZ 68. August 31-September 10. 

Euphilotes rita spaldingi (Barnes & McDunnough). Zuni Mts.: PoC, GS, 12 mi. S 
G, 4 mi. S FW (JS), M (JS), FW (TM, 1907). July 29-August 22. 

Euphilotes rita emmeli (Shields). Chuska Mts.: “E of Shiprock on dunes “ (JS in 
TM). August and September. 

Glaucopsyche piasus daunia (W. H. Edwards). Chuska Mts.: WP, WWC, BB, 3 
mi. NW Co, 8 mi. E WfL, RL, 12 mi. N WfL on TC, 2 mi. S WP. Zuni Mts.: 
2-3 mi. W M, PoC, GS, PT, FW (TM, 1910). May 16-June 30. 

Glaucopsyche lygdamus oro (Scudder). Chuska Mts.: WP, 2 mi. NW Toh, 1 mi. 
E WP, WWC, 2 mi. SE Co, 12 mi. N WfL on TC, 2 mi. S WP. Zuni Mts.: GS, 
PoC, PT, FW (TM, ?. Specimen is in Carnegie Museum. It may not be a Wood- 
gate specimen.). April 29-June 22. 

Lycaeides melissa pseudosamuelis Nabokov. Chuska Mts.: 2 mi. NW Toh, Toa, 2 
mi. SE Co, RL, Wf£L, WWC, BGR, L, 6 mi. SW Sn. Zuni Mts.: uBC, PoC, NDR, 
PrC, GS, CCT, FW (TM, 1907 & 1908). May 18—-August 20. 

Plebejus saepiolus whitmeri F. M. Brown. Chuska Mts.: 2 mi. S WP, ? TC (KR). 
June 17. 

Icaricia icarioides lycea (W. H. Edwards). Chuska Mts.: 3 mi. NW Toh, 2 mi. SE 
Co, 2 mi. W Toa, 1 mi. E WP, WfL, BG, 8 mi. N WfL on TC, OacTL, WhC, 
LP, 5 mi. NW Toa, 4 mi. N OS, 4 mi. S Co, BM. Zuni Mts.: GS, PT, IZC, PoC, 
5 mi. W M, DC, 3 mi. NE M, WT, GC, FW (TM, 1909 & 1910). May 23-August 
9. Bivoltine. 

Plebejus acmon texana Goodpasture. Chuska Mts.: 10 mi. N WP, 8 mi. SW Sn, 
RL, 16 mi. N WfL on TC, WEL, 2 mi. S WP, LP. Zuni Mts.: PoC, IZC, NMJ, 
FW (TM, 1910), BD (MT). May 1-+ August 19. 

Agriades rustica rustica (W. H. Edwards). Chuska Mts.: WP, RB, WWC, BG, 8- 
16 mi. N WfL on TC, 2 mi. NW Toh, 2 mi. S WP, LP. Zuni Mts.: PoC, DC. 
May 29-July 10. 

Apodemia mormo mormo (C. & R. Felder). Chuska Mts.: 6—8 mi. SW of Sn, 2 
mi. NW of Toh. Zuni Mts.: 3 mi. NW of NDR, FW (TM, 1907 & 1910). July 
24-September 18. At least bivoltine. Populations from the Rio Grande Valley, 
from Albuquerque south, are m. cythera (W. H. Edwards), m. mejicanus (Behr) 
or m. duryi (W. H. Edwards). Jemez Mts. populations I have personally seen 
are all m. mormo. Due to the great distances between known colonies in western 
New Mexico, we do not know where the dark DHW form (typical mormo) 
begins to replace the reddish form (cythera). Mixed or intermediate populations 
have not been found in the northern half of New Mexico. 

Apodemia nais (W. H. Edwards). Zuni Mts.: PoC, PrC, GS, FW (TM, 1910). June 


232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


24-August 15. Bivoltine. In New Mexico, nais feeds on Ceanothis and usually 
occurs in association with C. apama and E. pacuvius. 


LIBYTHEIDAE 


552b. Libytheana bachmanii larvata (Strecker). Chuska Mts.: 11 mi. E RoR. Zuni Mts.: 
DC, NMJ. May 27-August 19. 


NYMPHALIDAE 


562. Euptoieta claudia (Cramer). Records from vitually everywhere in NM. At least 
April—October. 

568. Speyeria nokomis nigrocaerulea (W. & T. Cockerell). I personally am convinced 
nigrocaerulea is a valid subspecies. Chuska Mts.: colonies in Apache Co., AZ, 
and San Juan Co., NM; numerous other dubious records; one authentic specimen 
from WWC (BG). Ova obtained from confined females from Apache Co. have 
been reared through on potted Viola without attempt by the larvae to enter the 
usual, troublesome Speyeria larval diapause. This procedure yields adults around 
1 Dec. (JJ). Zuni Mts.: BD (MT, sight record, 29 August 1967). At present, sheep 
and goats have rendered this locality incapable of supporting a nokomis colony. 
The WWC record in the Chuskas may represent a stray from an as yet unlocated 
colony. I fear the Zuni Mts. population is now extinct. July 20-August 10. This 
should be proposed as an endangered species. 

574f. Speyeria atlantis dorothea Moeck. Populations in Chuskas and on Mt. Taylor are 
definitely dorothea, not nikias (Ehrmann). Chuska Mts.: FAA, BG, RB, 1 mi. E 
WP, 8 mi. N WfL on TC, OacTL, 4 mi. SE Co, 8 mi. S ReR, 10 mi. WSW Sn. 
Zuni Mts.: ? no specific data (KR). June 22-July 27. 

592. Poladryas arachne (W. H. Edwards). Chuska Mts.: 8 mi. N WfL on TC, 2 mi. 
NW Toh, WWC, BG, Toa, 3 mi. NW Co, OacTL, RL, TL, BM, HP. Zuni Mts.: 
PoC, IZC, PrC, GC, NDR, LCC, FW (TM, 1907), BD (MT). May 14—-August 31. 
Bivoltine. 

597d. Thessalia leanira alma (Strecker). Chuska Mts.: 2 mi. NW Toh, WfL, L, RL, 
WhC, HP. Zuni Mts.: PoC, IZc, NMJ, CC, NDR, 6 mi. E R, FW (TM, 1907), 
BD (MT). May 17-August 9. 

599a. Chlosyne lacinia crocale (W. H. Edwards). Zuni Mts.: CG. August 15. 

609. Charidryas acastus (W. H. Edwards). Chuska Mts.: 2 mi. SE Co, WWC, Co, L, 
4.5 mi. S HP. Zuni Mts.: PoC, NMJ. May 13-June 15. 

623. Phyciodes tharos (Drury) ssp. Chuska Mts.: 3 mi. SE Co, 2 mi. NW Co, RB, 8 mi. 
W Toa. June 15-July 28. Unlike eastern populations and some which have be- 
come established around cultivated areas in southern NM, this large phenotype 
is univoltine. See Ferris and Brown (1981) for interesting remarks on the Chuska 
Mts. tharos. It seems possible that we are dealing with two species in New 
Mexico; one native, large, bright, mostly univoltine and found in undisturbed 
places; and another introduced from the East which is small, dark, multi-voltine 
and generally restricted to built-up areas. 

625c. Phyciodes pratensis camillus W. H. Edwards. Chuska Mts.: Toa, WWC, 2 mi. SE 
Co, L, WfL, RL, HP, 8 mi. N WfL on TC, 2 mi. NW Toh, WhC, 12 mi. N 
WEfL on TC. Zuni Mts.: 4 mi N NMJ, 2 mi. W M, PrC, SS, 5 mi. W R, BD, 
NDR, BV, PoC, GS, B, NMJ, IBC, FW (TM, ?). May 1—August 6. 

626b. Phyciodes picta canace W. H. Edwards. Chuska Mts.: BGR, 3 mi. W Sn, 6 mi. E 
Sn, WhC. May 12-+August 10. At least four broods. 

629b. Phyciodes mylitta nr. callina (Boisduval). Chuska Mts.: 1 mi. E WP, WP, 2 mi. 
NW Toh, RB, WWC, 8 mi. W Toa, 8 mi. E WfL, RL, 12 mi. N WfL on TC, 5 
mi. NW Toa. Zuni Mts.: BD, PoC, GS, PrC, IZC, uBC, FW. April 29-+July 3. 
callina (TL Sonora, Mexico) is not a very satisfying name for these populations. 
However, the name arizonensis Bauer is so vaguely described as to make one 
unsure as to what it is applied. 


VOLUME 38, NUMBER 3 233 


631a. 


631g. 


638a. 


642. 


647a. 


648a. 


649b. 


650. 


651. 


652. 


653a. 


656. 


665e. 


668a. 


717Ta. 


729. 


738a. 


734a. 


7395a. 


748a. 


Occidryas anicia alena (Barnes & Benjamin). Chuska Mts.: 2 mi. SE Co, N slope 
BG, Co, L, 1 mi. NW Toh, W£L, 2 mi. SW Co, 12 mi. S Toa, 4.5 mi. S HP, 5 
mi. E G. Zuni Mts.: GS, NDR, NMJ. April 30-June 15. 

Occidryas anicia chuskae Ferris and R. Holland. Chuska Mts.: RB, 8 mi. N WfL 
on TC, 8 mi. W Toa, OacTL (CDF & RH), 10 mi. E of WfL on WIC, 6 mi. 
NW Toa. June 22—+July 8. Separated from alena temporally (latest alena June 
15, peak in May) and altitudinally (alena 5500’ to 7000’ in Chuskas, chuskae 
7300’ to 9000’). 

Polygonia satyrus satyrus (W. H. Edwards). Chuska Mts.: Toa, 8 mi. W Toh. Zuni 
Mts.: NDR. April 21-+July 11. 

Polygonia zephyrus (W. H. Edwards). Chuska Mts.: BG, RB, FAA, 3 mi. NW 
Toh, WWC, WP, 2 mi. S WP. Zuni Mts.: DC, 4 mi. S FW. April 23-+ August 
10. 

Nymphalis californica californica (Boisduval). Chuska Mts.: 2 mi. SE Co, 4 mi. 
SE Co, BG, Toa. Zuni Mts.: PoC, DC. April 21-July 21. 

Nymphalis antiopa antiopa (Linnaeus). Common everywhere in NM at interme- 
diate elevations (Upper Sonoran and Canadian zones) at least April through 
October. 

Aglais milberti furcillata (Say). Chuska Mts.: ? no specific data (KR). This species 
is found on other mountains in northwestern NM. 

Vanessa virginiensis (Drury). Chuska Mts.: Toa, 8 mi. W Toa, 264. Zuni Mts.: 3 
mi. W M, WT, DC, NJ. May 11-+July 22. 

Vanessa cardui (Linnaeus). Abundant everywhere and every year in NM from 
April to November. Migrations are much heavier in some years than others. In 
years of heavy migration, a few specimens always may be taken several weeks 
before the main migration arrives. Occasionally during migrations, the popula- 
tion is so dense as to interfere with nighttime light trapping of moths. 

Vanessa annabella (Field). Chuska Mts.: WP, 2 mi. SE Cove, RB, Toh, WfL, TL. 
Zuni Mts.: IBC, NDR. May 19-—+June 21. 

Vanessa atalanta rubria (Fruhstorfer). Chuska Mts.: WfL, 2 mi. SE Co, Toa, 11 
mi. E RoR, 8 mi. N WfL on TC, 264. Zuni Mts.: NDR, DC, NMJ, OjR (RB). 
May 11-+July 11. Population may have a migratory component, as it is much 
more common in years of strong cardui migrations. 

Junonia coenia Hubner. Chuska Mts.: RL, WhC, Toa, 12 mi. E RoR. July 1- 
August 6. All Chuska Mts. records from 1978, when a migration occurred. Not 
frequently encountered in the northwestern quarter of NM, although taken in 
numbers in the Jemez Mts. in 1983 and 1984. 

Basilarchia weidemeyerii angustifascia Barnes and Benjamin. Chuska Mts.: 2 mi. 
SE Co, WWC, BG, 4 mi. E WIL, 2 mi. E CP, BM, 2 mi. SW Co, LP. Zuni Mts.: 
4 mi. N NMJ, NDR, GS, PoC, PrC, MRC, FW (TM, 1909). May 30-+July 21. 

Adelpha bredowii eulalia (Doubleday & Hewiston). Chuska Mts.: 8 mi. N WfL 
on TC, WEL, 6 mi. W Sn, WWC. Zuni Mts.: GS, PoC. May 28-September 2. 


SATYRIDAE 


Cyllopsis pertepida dorothea (Nabobov). Chuska Mts.: 2 mi. SE Co, Toa, 2 mi. 
NW Toh, HP. Zuni Mts.: PoC, IZC, GS, NDR, LCC, PRC, MC, FW (TM, ?), 
BD (MT). June 18—-August 18. 

Coenonympha ochracea W. H. Edwards. Chuska Mts.: ? BG (KR). 

Cercyonis meadii meadii (W. H. Edwards). Chuska Mts.: 6 mi. W Sn, Toa, 2 mi. 
SE Co, 8 mi. SW Sn. July 14-September 2. C. meadii and sthenele masoni 
intergrade in the Chuskas. 

Cercyonis sthenele masoni Cross. Chuska Mts.: 1 mi. SE Co, 6 mi. W Sn, Toa, 8 
mi. SW Sn, 8 mi. NW Sn. July 13-September 2. 

Cercyonis oetus charon (W. H. Edwards). Chuska Mts.: RB, 5 mi. S RB, 8 mi. W 
Toa, 8 mi. N WfL on TC, 3 mi. W Toa, 12 mi. N WfL on TC, 6 mi. W Sn, 
OacTL, 4 mi. E WfL, 10 mi. E WfL on WfC, BM. June 22—August 5. 

Neominois ridingsii. Zuni Mts.: 1ZC, uBC, 1 mi. N FW. May 31l-June 27. 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


DANAIDAE 


760. Danaus plexippus (Linnaeus). Chuska Mts.: 1 mi. W Toa, 8 mi. N WfL on TC, 
LP, HP. Zuni Mts.: NDR, FW. May-September. 

76l1b. Danaus gilippus strigosus (Bates). Chuska Mts.: Toa, 11 mi. E RoR. Zuni Mts.: 
NDR, MRC. May 27-July 22. 


ACKNOWLEDGMENTS 


The author is indebted to R. E. Stanford, C. D. Ferris and L. D. Miller for reviewing 
this article. The Toliver manuscript is an indispensable reference for faunal studies of 
New Mexico. It is most regrettable it was never published. 


LITERATURE CITED 


BROWN, F. M. 1983. The type of Argynnis apacheana Skinner. J. Lepid. Soc. 37:79- 
80. ) 

, with D. Err & B. ROTGER. 1957. Colorado Butterflies. Denver Museum of 

Natural History, Denver. viii + 368 pp. 

& L. D. MILLER 1980. The types of the Hesperiid butterflies named by William 

Henry Edwards Part II, Hesperiidae: Hesperiinae, Section II. Trans. Amer. Entomol. 

Soc. 106:43-88. 

& P. A. OPLER. 1970. The types of the Lycaenid butterflies described by 
William Henry Edwards. Trans. Amer. Entomol. Soc. 96:19-77. 

CHANG, V. C. S. 1963. Quantitative analysis of certain wing and genitalia characters 
of Pieris in western North America. J. Res. Lepid. 2:97-125. 

FERRIS, C. D. 1976. A checklist of the butterflies of Grant County, New Mexico and 
vicinity. J. Lepid. Soc. 30:38—49. 

& F. M. BRowN. 1981. Butterflies of the Rocky Mountain States. University 
of Oklahoma Press, Norman. xviii + 442 pp. 

HOLLAND, R. 1974. Butterflies of six central New Mexico mountains, with notes on 
Callophrys (Sandia) macfarlandi (Lycaenidae). J. Lepid. Soc. 28:38—-52. 

KEARNEY, T. H. & R. H. PEEBLES. 1964. Arizona Flora. University of California Press, 
Berkeley. viii + 1085 pp. 

MILLER, L. D. & F. M. Brown. 1981. A Catalogue/Checklist of the Butterflies of 
America North of Mexico. Lepid. Soc. Mem. 2. vii + 280 pp. 

PEARCE, T. M., with I. S. Cassipy & H. M. PEARCE. 1975. New Mexico Place Names, 
A Geographical Dictionary. The University of New Mexico Press, Albuquerque. xvi 
+ 187 pp. 

SKINNER, H. 1911. A new variety of Megathymus yuccae (Lepid.). Entomol. News. 
22:300. 

TOLIVER, M. E. 1978. Distribution of butterflies (Lepidoptera: Hesperioidea and Pa- 
pilionoidea) in New Mexico. (Unpublished. ) 

WILLIAMS, R. C. 1914. One hundred butterflies from the Jamez [sic] Mountains, New 
Mexico (Lepid.), with notes and descriptions of a new species. Entomol. News 25: 
263-268. 














Journal of the Lepidopterists’ Society 
88(3), 1984, 235 


GENERAL NOTES 


THE LARCH CASEBEARER, COLEOPHORA LARICELLA (HUBNER) 
(COLEOPHORIDAE), IN WESTERN WASHINGTON 


The larch casebearer, Coleophora laricella (Hiibner), is a Palearctic moth which was 
first reported from the Pacific Northwest in 1957 by Denton (1958, U.S. For. Serv. Res. 
Note 51:1-6). Denton and Tunnock (1971, U.S.D.A. For. Pest Leaflet 96: fig. 1) mapped 
the species’ range in the northwestern United States and adjacent paris of Canada. Since 
1971, field parties from the University of Washington, Seattle, have observed larvae on 
western larch, Larix occidentalis Nuttall, from two localities on the east slope of the 
Washington Cascades Range: Chelan Co., 12.5 km SW Leavenworth, Bridge Creek 
Campground; and Kittitas Co., 8.8 km SE Cle Elum, Elk Heights. In March 1981, I 
found third instar larvae feeding on the new foliage of a European larch, L. decidua 
Miller in King Co., Seattle, Univ. Washington campus. Individuals were subsequently 
reared and voucher specimens deposited in the collection of the University of Washington 
College of Forest Resources. In the spring of 1981 and 1982, infestations of this moth 
were found on European larch at Green Lake, about four kilometers northwest of the 
campus. 

Mr. Richard Johnsey, State Forest Entomologist, Washington Department of Natural 
Resources, who maintains western Washington records of pest insects, informed me that 
C. laricella had not been previously reported west of the Cascades in this state. How the 
moth crossed the Cascades (the lowest pass is 922 m) is conjectural. Prevalent winds are 
normally from west to east or north to south. The species may have been transported 
with nursery stock, or its spread westward may be natural. 

I thank Dr. Robert Gara, University of Washington, and Dr. Frederick H. Rindge, 
American Museum of Natural History, for their encouragement and critical commentary 
on the manuscript. 


SANFORD R. LEFFLER, College of Forest Resources, University of Washington, Se- 
attle, Washington 98195. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 235-236 


THE GESNERIACEAE AND BIGNONIACEAE AS FOOD-PLANTS 
OF THE LEPIDOPTERA 


Robert K. Robbins and Annette Aiello in their paper, Foodplant and Oviposition 
Records for Panamanian Lycaenidae and Riodinidae (1982, J. Lepid. Soc., 36(2):65- 
75), with their single record of a gesneriad as a lepidopterous food-plant and their 
quotation from Ehrlich and Raven’s 1964 paper, Butterflies and plants, a study in co- 
evolution, that plants belonging to the Gesneriaceae, Bignoniaceae and Begoniaceae are 
not used or are under-utilized as lepidopterous food-plants, reminded me of a recent 
experience. 

We have in East Africa a number of indigenous Gesneriaceae, including the wild 
ancestor of the very popular African Violet, or Saintpaulia, as well as numerous intro- 
duced species grown as pot plants in greenhouses and open verandahs, but I have only 
recently obtained a record of a gesneriad being eaten by a lepidopterous larva. On two 
separate occasions larvae of the polyphagous sphingid, Coelonia mauritii Btlr., have 
been found feeding on Aeschynanthus marmoratus, an introduced cultivar from Thai- 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


land, growing in a hanging basket suspended from the roof of an open-sided orchid house 
in Mombasa. 

The following food-plants have previously been recorded for C. mauritii: Acanthus 
(Acanthaceae), Cissus (Ampelidaceae), Bignonia, Fernandoa magnifica, Markhamia 
platycalyx, Millingtonia hortensis, Newbouldia imperialis, Spathodea, Tecoma, Teco- 
maria (Bignoniaceae), Cordia (Boraginaceae), Dahlia (Compositae), Convolvulus, Ipo- 
maea (Convolvulaceae), Coleus, Pycnostachys, Salvia (Labiatae), Buddleia, Lachnopylis 
(Loganiaceae), Jasminum (Oleaceae), Lycopersicum, Nicotiana, Solanum (Solanaceae), 
Hebe speciosa (Scrophulariaceae), Clerodendron, Duranta, Lantana, Stachytarpheta 
indica (Verbenaceae). 

Contrary to Ehrlich and Raven’s comment, the Bignoniaceae do provide food-plants 
for a considerable number of Lepidoptera, mostly Heterocera it is true, in East Africa, 
but I have far fewer records for India. Below is a complete list of my records: 


Bignonia—Spilosoma investigatorum Karsch (Arctiidae), Acherontia atropos L., Coe- 
lonia mauritii Btlr. (Sphingidae); Fernandoa—Cymothoe coranus Gr. Sm. (Nymphali- 
dae), Epiphora mythimnia Westw. (Saturniidae), C. mauritii (Sphingidae), Mazuca 
strigicincta Wlk. (Noctuidae), Hyblaea euryzona Prout (Pyralidae); Jacaranda—Pachy- 
pasa sericeofasciata Auriv. (Lasiocampidae); Kigelia—C. coranus, Asterope boisduvali 
Wllgrn. (Nymphalidae), Mussidia nigrivenella Rag., M. fiorii T. & deJ., Zebronia phe- 
nice Cr., Udea ablactalis Wlk. (Pyralidae); Podranea—A. atropos (Sphingidae); Mark- 
hamia—Euproctis molundiana Auriv. (Lymantriidae), Pachypasa subfascia Wlk., Pseu- 
dometa castanea Hamps. (Lasiocampidae), Phiala atomaria Holl. (Eupterotidae), C. 
mauritii, Macropoliana natalensis Btlr., Andriasa contraria Wlk. (Sphingidae), Pera- 
todonta olivaceae Gaede (Notodontidae), Latoia chapmani Kirby, L. hexamitobalia Tams, 
L. vivida Wlk., L. urda Druce, L. viridicosta Hamps. (Limacodidae), Salagena atridisca 
Hamps. (Metarbelidae), Lycophotia ablactalis Wlk., M. strigicincta (Noctuidae), Com- 
ibaena leucospilata Wlk. (Geometridae), Hyblaea puera Cr., H. euryzona, Polygram- 
modes junctilinealis Hamps., Z. phenice, Pyrausta fulvilinealis Hamps. (Pyralidae); 
Millingtonia—Hypolycaena philippus F. (Lycaenidae), A. atropos, C. mauritii, Pemba 
favillacea W\k. (Sphingidae); Newbouldia—Argyrostagma niobe Weym. (Lymantri- 
idae), Agrius convolvuli L., C. mauritii, A. contraria (Sphingidae); Spathodea—Holocera 
smilax Ang. (Saturniidae), A. atropos, C. mauritii, M. natalensis, Poliana witgensis Strd., 
A. contraria, Cephonodes hylas L., Hippotion osiris Dalm. (Sphingidae), Z. phenice 
(Pyralidae); Stereospermum—P. subfascia (Lasiocampidae), Z. phenice (Pyralidae); Te- 
coma—A. atropos, C. mauritii (Sphingidae), U. ablactalis (Pyralidae); Tecomaria— 
Spilosoma lutescens Wlk. (Arctiidae), A. atropos, C. mauritii (Sphingidae). My Indian 
records are the sphingids Acherontia styx Westw., A. lachesis F., and Psilogramma 
menephron Cr. on species of Tecoma, Stereospermum, Bignonia and Spathodea; pre- 
sumably Hyblaea puera also feeds on Bignoniaceae in India, but I have no records. 

The Begoniaceae is another story, the only East African record I have is Bracharoa 
quadripunctata Wllgrn. (Lymantriidae) on Begonia sp., and for India the sphingids 
Theretra clotho Drury, T. latreillei Macleay and Rhyncholaba acteus Cr., also on Be- 
gonia spp. 

I cannot help feeling that Ehrlich and Raven would have come to some very different 
conclusions if they had included the Heterocera in their survey. 


D. G. SEVASTOPULO, F.R.E.S., P.O. Box 95617, Mombasa (Nyali), Kenya. 


VOLUME 38, NUMBER 3 Jal 


Journal of the Lepidopterists’ Society 
38(3), 1984, 237-242 


NOTES ON THE NATURAL HISTORY OF PAPILIO VICTORINUS 
DOUBL. (PAPILIONIDAE) IN NORTHEASTERN COSTA RICA 


Papilio victorinus Doubl. (Papilionidae) is a member of the “homerus group” of 
“fluted” swallowtail butterflies inhabiting Central America and Mexico (Seitz, 1924, Mac- 
rolepidoptera of the World, Vol. 5, Kernan, Stuttgart). The caterpillar (instar not men- 
tioned) and pupa were described by Schaus (1884, Reise Novara, Lepid., Papilio 4:101) 
from Mexico. All early stages and a larval food plant were reported for P. victorinus 
from El Salvador by Muyshondt et al. (1976, Rev. Soc. Mex. Lepid. 2:77-90). One 
specimen of this species in the pinned collection of the Costa Rican National Museum 
bears a label stating “reared 26 May 1979 on Persea americana.’ Muyshondt et al. (op. 
cit.) also report Persea (Lauraceae) as the larval food plant of this butterfly. Butterflies 
of the “homerus group” are known to feed as caterpillars on several plant families, most 
notably Lauraceae, Hernandiaceae, Rubiaceae, Malvaceae, and Convolvulaceae (Scriber, 
unpubl. manuscript, Latitudinal gradients in larval feeding specialization of the world 
Papilionidae (Lepidoptera)—-A supplementary table of data, for Psyche 80:355-373). 
Herein, I report for the first time the purported feeding association of P. victorinus with 
Hernandiaceae in northeastern Costa Rica, a discovery not unexpected given the known 
larval food plant associations of the “homerus group’ species (Scriber, op. cit.). I also 
provide further documentation of the early stages to supplement those of Muyshondt et 
al. (op. cit.) for El Salvadoran populations. 

At 1145 h on 2 February 1977, I observed a large black swallowtail butterfly place a 
total of four eggs on a leafy tree sapling (about 1.5 m tall) in a partly shaded clearing 
within mixed primary and secondary “premontane tropical wet forest” at “Finca La 
Tigra,’ near La Virgen (10°23’N, 84°07'W; 220 m elev.), Sarapiqui District, Heredia 
Province. This butterfly had large diffuse areas of bluish green on the upper surfaces of 
the hind wings. Although I could not readily determine the species, judging from my 
experience with observing other swallowtail butterflies in Costa Rica over the past sixteen 
years, I ruled out familiar species such as Battus belus varus Kollar, B. polydamas 
Linnaeus, B. crassus lepidus Cramer, Eurytides pausanias prasinus Roth. & Jordan, and 
Papilio cleotas archytas Hoppfer. In an earlier draft of this paper, I erroneously identified 
the butterfly in question as P. birchalli Hew. But after reading the reviewer’s comments 
and rechecking descriptions of this species and consulting further the Muyshondt et al. 
(op. cit.) reference on P. victorinus early stages, I am assuming my species to be P. 
victorinus. One source of confusion was examining a wild-caught female specimen in 
the Costa Rican National Museum labeled as both P. victorinus and P. birchalli. The 
occurrence of P. birchalli in Costa Rica is questionable (e.g., Seitz, op. cit.). Because I 
was unable to rear the four eggs through to adulthood, I can only state that the species 
in question is purportedly P. victorinus. Further indirect evidence against it being P. 
birchalli, for example, is the systematic placement of this species within another “fluted” 
swallowtail group, the “scamander group,” whose species with known life cycles are not 
associated with Hernandiaceae (Scriber, op. cit.). Given these considerations, I am assum- 
ing the species to be P. victorinus. Owing to the fact that it is often very difficult to 
obtain oviposition records for Papilio species in the wild (J. M. Scriber, pers. comm.), 
incomplete rearing data such as mine in this particular instance do provide an initial 
observation on which to build further studies, even though it may take many more years 
before myself or another researcher witnesses a large black Papilio placing eggs on 
Hernandiaceae in northeastern Costa Rica. 

During oviposition, the butterfly in question made several swooping low passes over 
the small tree, each time placing an egg on the shaded forest plant (Fig. 1). The first 
honey-colored spherical egg (2 mm dia.) was placed on the ventral surface of a mature 
leaf. A second egg was placed, seconds later, on the upper surface of the same leaf. Prior 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


~ 


Fic. 1. Egg-placement forest habitat (above) and larval food plant (below, broad- 
leaf plant immediately to the left of the insect net) of Papilio birchalli Hew. (Papilion- 
idae) at “Finca La Tigra’ in northeastern Costa Rica. 


to ovipositing a third egg, the butterfly flew swiftly into the upper reaches of the forest, 
and then returned to place another egg on the ventral surface of a meristem leaf. A 
fourth egg was quickly placed on the ventral surface of the same leaf. Within approxi- 
mately four minutes, the insect placed four eggs on two different leaves of the tree, 





VOLUME 38, NUMBER 3 239 


apparently exhibiting a lack of “preference” for meristem versus mature leaf surfaces as 
oviposition cues. I left one egg on the plant and collected the other three for rearing. 

Before leaving the site when collecting the eggs, I marked the tree with a small yellow 
tag (plastic) secured with copper wire. I marked the tree in order to make subsequent 
observations for additional eggs and caterpillars of this Papilio over the next several 
years. In doing so I also ensured accurate food plant voucher specimens for confirming 
identification of the plant. At the time the oviposition was observed, the tree had no 
flowers or fruits, rendering it difficult to make positive determination of the plant. A 
voucher of fresh leaves was collected at this time for determination, and over the follow- 
ing seven years, three additional vouchers were taken for determinations. 

Based upon the examination of fresh fragmentary material collected from the tree for 
the first three times, three different well-known botanists familar with the Costa Rican 
flora independently determined the plant to be in the Araliaceae. Based upon a review 
of the manuscript when previously submitted to this journal and in which the food plant 
determination was challenged, I collected the fourth and final voucher from the tree (2 
August 1984) and arranged for one of the botanists, Luis Diego Gomez, to re-examine 
the material. In conferring with another botanist, Luis Fournier, it was determined that 
the plant in question was Hernandiaceae, either Hernandia sonora or H. guianensis (L. 
D. Gomez, letter to A. M. Young, 3 August 1984). Mr. Gomez indicated to me that 
several features of the material lead one to believe that the plant is Araliaceae. Thus he 
writes: “The different lengths of the petioles, lustrous leathery leaves and the methylated 
aroma of crushed leaves, suggested an aralia.”’ The methylated compounds underlying 
aroma of the crushed leaves are flavonoids also found in the Araliaceae and Umbelliferae 
(L. D. Gomez, pers. comm.). This distinctive aromatic property is also encountered in 
the Lauraceae, the other known larval food plant of P. victorinus and the “homerus 
group (e.g., Muyshondt et al., op. cit.). In Costa Rica, the Hernandia in question (Fig. 
1) is locally called “aguacatillo” (small avocado, little avocado), as Gomez conveyed to 
me, not only because the twigs faintly resemble those of Lauraceae but because of the 
aromatic bark and leaves.” - 

The eggs were kept in a large, clear, plastic bag maintained tightly shut and containing 
fresh cuttings from the food plant. The honey-colored egg (Fig. 2) darkened noticeably 
a day before hatching, and hatching occurred in eight days. The first instar larva (Fig. 
2) immediately devoured the entire empty egg shell and readily everted a reddish orange 
osmeterium at the slightest provocation. The first instar larva is 6 mm long at the time 
of hatching, bears a glossy, smooth dark-brown head capsule, a dorsal pair of long, orange 
tubercles on the first thoracic segment; a second pair of short, dark brown tubercles are 
borne laterally on this segment. The second thoracic segment has one pair of short, dark 
brown tubercles and a lateral long pair (also brownish). Tubercles with short brown or 
black setae. The same pattern of two pairs of tubercles occurs on the third thoracic 
segment. The first three abdominal segments also have two pairs each of much shorter 
brownish tubercles; those of the fourth are white. Segments 5-8 with tubercles as seg- 
ments 1-3 of the abdomen. Segments 9-10 with only dorsal, long, white tubercles. The 
elongate tubercles of both the first thoracic segment and the final abdominal segments 
give the body an illusory “bi-forked’”’ appearance. The anal plate is dark brown; the 
remainder of the body is a patchwork of brown and white blotches. 

Second instar larva similar in appearance to the first. Third instar (Fig. 8) without the 
prominent tubercles of the previous instars and now with the anterior third of the body 
greatly “swollen” in appearance. Resembles a typical P. cresphontes (Cramer) third 
instar. 

Lervae perch on individual silken mats on dorsal surfaces of leaves, both in the lab- 
oratory and as observed for the one caterpillar left to hatch in the wild; quickly rears up 
first % of body when disturbed, and holds this position for a few minutes. The “wild” 
caterpillar disappeared as a third instar on 15 February 1977, fifteen days after the egg 
was placed there. By this time the caterpillar had moved off the original leaf where the 
egg had been placed. 

Fourth instar larva very similar to third instar, but with row of roundish blue spots 
laterally on the abdominal region. Mimics of fresh bird dropping, as seen in the cater- 


240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Egg (upper left), a first instar (upper right and lower left), and early second 
instar (lower right) larva of P. birchalli. The photograph of the second instar was taken 
directly in the wild, on the food plant (Araliaceae, prob. Dendropanax sp.); note the 
silken mat the larva rests upon, in sunlight on upper side of a leaf. 


pillars of many Papilio species. The fifth instar larva (Fig. 3) is very differently colored 
than the previous instars, becoming patterned in shades of green and brown and attaining 
a body length of about 55 mm (n = 8) in about 10 days. The appearance of the fifth 
instar in my study is virtually identical to that of Muyshondt et al. (op. cit.) for this 


VOLUME 38, NUMBER 3 241 


y XX 
N 


- 


y ~ wes ; 2 
ee, ' : SS | 





Fic. 3. Third instar larvae (above, in laboratory culture), and fifth instar larva (be- 
low, left) of P. birchalli. Also shown is the larval food plant individual with apical section 
removed for determination studies. 


species in El] Salvador. The “‘cross-like” dorsal trunk pattern (Fig. 3) may be typical for 
species in the “homerus group.’ Owing to an absence of additional food plant material, 
all three caterpillars died prior to pupation but were probably very close to pupation. 
For the time of study, the larval period was 45 days at about 25—29°C. 

Although apical sections of the food plant individual were removed for identification 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


purposes later in the study (Fig. 3), the plant grew to a height of about 3.0 m by 
December 1982, and average leaf size at this time was about half of that at the time 
oviposition was observed. No other eggs or caterpillars of P. victorinus were found on 
the tree over the following seven years (with about a total of 26 days per year at three 
different times per year for examining the tree). 

Papilio species within the “scamander” and “homerus” groups appear to be typified 
as Magnoliales- and Laurales-feeders (Scriber, op. cit.), and as exemplified by the asso- 
ciation of P. victorinus with Persea in both El Salvador (Muyshondt et al., op. cit.) and 
Costa Rica (Costa Rican National Museum specimen label data) as well as with Hernan- 
dia (this report). Both the Lauraceae and Hernandiaceae fall within the Laurales (Cron- 
quist, 1981, An Integrated System of Classification of Flowering Plants, Columbia Univ. 
Press, New York). The similarity of aromatic properties of freshly crushed leaves in both 
groups, namely flavonoids (L. D. Gomez, pers. comm.), suggests a common ovipositional 
cue for P. victorinus. Yet, since these methylated compounds are also found in the 
Araliaceae and Umbelliferae, other known Papilio larval food plant groups (Scriber, op. 
cit.), P. victorinus must cue into still other factors in the food plant selection process, 
rendering the insect an excellent phytochemist-. 

I sincerely thank Luis Diego Gomez, Luis Jorge Poveda, and Gary S. Hartshorn for 
making initial determinations of the larval food plant, and to an anonymous reviewer 
who most admirably took the time to check carefully these determinations based upon 
the plates submitted with an earlier version of the manuscript. A special thanks to Luis 
Diego Gomez for taking the time in August 1984 to confer further with me on the plant 
identification, and to Dr. Luis Fournier for his assistance as well. And to whoever placed 
the “P. birchalli” name label on the female P. victorinus specimen (one of two specimens 
in the collection as of 17 August 1984) at the Costa Rican National Museum, please check 
it since it is a source of confusion with identification of the species. I thank Dr. J. Mark 
Scriber for reading the earlier draft and for helpful discussions which ensued from the 
editorial process. In the latter context, I also thank Dr. Thomas D. Eichlin, Journal Editor. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 532838. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 242-245 


“EDGE EFFECT” IN OVIPOSITION BEHAVIOR: A NATURAL 
EXPERIMENT WITH EUCHLOE AUSONIDES (PIERIDAE) 


The “edge effect,” whereby isolated host plant individuals tend to receive dispropor- 
tionate egg loads, has been documented in a variety of insects, and several authors have 
commented recently on mechanisms to account for it in butterflies. These mechanisms 
may be arranged in a proximate-ultimate causal hierarchy and may not be mutually 
exclusive, but attention has focused primarily on whether the “edge effect” is an adaptive 
characteristic produced by natural selection, or essentially a statistical artifact with no 
evolutionary significance (Shapiro, 1981, Am. Nat. 117:276-294; Courtney & Courtney, 
1982, Ecol. Entomol. 7:131-137; Mackay & Singer, Ecol. Entomol. 7:299-303). 

Another phenomenon affecting egg dispersion in various insects, including butterflies, 
is “egg-load assessment,” wherein ovipositing females react positively or negatively to 
the presence of previously laid, usually highly conspicuous, eggs (for butterflies see Raush- 
er, 1979, Anim. Beh. 27:1034-1040; Shapiro, 1980, J. Lepid. Soc. 34:307-315; Shapiro, 
1981, Am. Nat. 117:276-294; Singer & Mandracchia, 1982, Ecol. Entomol: 7:327-330). 
The interactions of these two phenomena may be complex and difficult to interpret in 
analyzing field egg-dispersion data. 


VOLUME 38, NUMBER 3 243 


TABLE 1. Distribution of Euchloe ausonides eggs on Brassica inflorescences at Suisun 
City, California, 28 March 1983. 


Number of jnfloxescenees bearing: Red eggs Green eggs 


8 1 
14? 0 
3 1 
6 0 
1 0 
3 0 
19 0 
Totals: 54 (35 with eggs) ll 44 
Mean red eggs/inflorescence having only red eggs: 1.00. 
Mean red eggs/inflorescence having both red and green eggs: 1.00. 
Mean green eggs/inflorescence having only ate green eggs: 1.71. 


Mean green eggs/inflorescence having bo green eggs: 1.00. 
2 includes 1 egg laid on adjacent leaf (Fig. ee 


OonRWONEFE © 


The large marble, Euchloe ausonides Lucas, is a member of the red-egg, inflorescence/ 
infructescence-feeding pierid guild in western North America, and engages in egg-load 
assessment (Shapiro, 1981, op. cit.): the mean number of eggs/inflorescence bearing any 
eggs is normally almost exactly unity. During the winter of 1982-83 in northern Cali- 
fornia, rainfall totals generally exceeded 175% of 30-year norms, and 200% was not 
unusual. At Suisun City, Solano County, where I have studied a population of E. auson- 
ides since 1978, much of the breeding habitat was inundated from 4-11 weeks to a depth 
of 15-80 cm. This unusual situation permitted a test of the flexibility of oviposition 
behavior, given a drastic shortage of sites: would the characteristically even dispersion 
of eggs change as “edge effect” became more important than “egg-load assessment’? 

Host plants—weedy mustards of the genus Brassica—are normally abundant at Suisun. 
By late March, from two to four species may be in a suitable phenophase (flower buds 
present) for oviposition by E. ausonides to occur. Between 1973 and 1981, the first flight 
was between 6-16 March at this site, oviposition commencing almost at once. (1982 
populations were so sparse that the dates may not be reliable.) In 1983 the first males 
were seen on 26 March. By 28 March both sexes were common, and an egg census was 
done. Every Brassica plant in a 1.5 ha field was examined thoroughly. Normally this 
would be impossible—there would be many thousands—but on this occasion only 140 
plants could be found. Of these only 22 had any inflorescences judged suitable for ovi- 
position—a total of 54 of them. At least three females were observed ovipositing on the 
site on 28 March. The distribution of eggs on the 54 inflorescences is given in Table 1. 

More than a third of the inflorescences bearing eggs bore more than one. Euchloe 
ausonides eggs are green when laid, changing to red by the next day. For green (same 
day) eggs only, the mean number of eggs/inflorescence bearing eggs was 1.71. Before 
28 March I had never seen a four-egg inflorescence, but on that day I found three. One 
female was seen laying on an inflorescence known to bear one green egg. One egg was 
found on an upper leaf adjacent to a very rudimentary inflorescence, too small to permit 
the female to balance upon it while laying. This is the first E. ausonides egg I have ever 
found on a leaf. Fig. 1 illustrates some of these situations. 

The distribution of the eggs among Brassica species is of interest. All four known hosts 
at the site were present, but their phenologies differ enough that their reactions to the 
flood were quite different. B. campestris L., the first to germinate and bloom, was 
represented by only three individuals—all past bloom, bearing green fruit and no eggs. 
B. Kaber (DC.) Wheeler, on the average somewhat later, had been harmed most. Six 
individuals were present, all in flower; three of these had usable buds, and all the sus- 
ceptible inflorescences received multiple ovipositions. B. nigra (L.) Koch, last of the 
annual mustards to bloom, had germinated as the flood receded and was still mainly 
vegetative. It was the commonest species, but few plants had well-defined buds—the one 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Brassica inflorescences collected at Suisun City, California on 28 March 1983 
showing unusual ovipositions by Euchloe ausonides. Upper left: B. Kaber with one red 
and one green egg. Upper right: B. Kaber with four green eggs. Lower left: B. geniculata 
with four green eggs. Lower right: B. nigra with egg laid on cauline leaf adjacent to 
rudimentary buds. (Photos by Samuel W. Woo.) 


illustrated, with an egg on a leaf, was one of the two most advanced—and very few eggs 
were found on them. B. geniculata (Desf.) J. Ball, a short-lived perennial, had survived 
the flood and bolted. It was in prime condition for oviposition—early flowering, with 
many buds—and received most of the eggs. The most important criterion for oviposition 
was obviously phenophase and not species or size. 

Previous studies (Shapiro, 1981, op. cit.) have shown that newly laid, green eggs are 


VOLUME 38, NUMBER 3 245 


not deterrent to females in the red-egg guild and have failed to support the existence of 
an oviposition-deterrent pheromone. This “natural experiment’’ supports these conclu- 
sions. The greatly increased incidence of multiple oviposition suggests that when host 
density is reduced by 3-4 orders of magnitude while population density is normal, the 
entire stand of hosts may demonstrate “edge effect” —at least early in the flight, when 
most eggs are green. Theoretically, as the flight proceeds, the presence of more red eggs 
should deter multiple ovipositions and perhaps encourage female dispersal. Unfortu- 
nately, it was not practical to test this prediction, given the rate of turnover of inflores- 
cences and the rapid maturation of the many B. nigra at the Suisun site. The ability of 
“edge effect” to dominate the pattern of egg dispersion in this unusual situation, however, 
does tend to confirm that “edge effect” is a statistical consequence of female behavior; 
it does not clarify the evolutionary origin of that behavior. 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 245 


EPIBLEMA LUCTUOSANA A. BLANCHARD, A HOMONYM, IS 
CHANGED TO EPIBLEMA LUCTUOSISSIMA, NEW NAME 


From Dr. Leif Aarvick (Tarnveien 6, N-1430 As, Norway), I received the following 
information, for which I thank him very much: “Blanchard describes a species which he 
calls Epiblema luctuosana. Unfortunately there is another Epiblema luctuosana in Eu- 
rope (E. luctuosana Duponchel, which is a synonym of E. scutulana Den. & Schiff). 
Thus luctuosana A. Blanchard is a homonym.”’ 

I propose to change the name of the species I described as E. luctuosana (1979, J. 
Lepid. Soc. 33(3):184) to Epiblema luctuosissima A. Blanchard. 


ANDRE BLANCHARD, 3023 Underwood St., Houston, Texas 77025. 


Journal of the Lepidopterists’ Society 
88(3), 1984, 245-249 


SCHIZURA RUSTICA (SCHAUS), A NOTODONTID MOTH DEFOLIATING 
HERRANIA AND THEOBROMA SPECIES (STERCULIACEAE) 
IN COSTA RICA 


Herein, I report for the first time the association of the “medium-sized” (approx. 37 
mm spread wingspan), dull brown and mottled gray notodontid moth Schizura rustica 
(Schaus), with Herrania albiflora Goudot (Sterculiaceae) as a larval food plant at one 
locality in Costa Rica and the acceptability of the closely related Theobroma cacao L. 
(also Sterculiaceae) as an alternate food plant. My report includes observations on the 
role of this moth as a serious defoliator of H. albiflora as well as offering some preliminary 
autecological and natural history notes on the life cycle and larval feeding behavior. 
Although much information has accumulated over the years on the insect herbivores 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


associated with T. cacao throughout the tropical regions of the world (e.g., Entwistle, P. 
F., 1972, Pests of Cocoa, Longman, London; Saunders, 1979, Plagas Insectiles De America 
Central, Turrialba, Costa Rica: CATIE). Given the very close evolutionary affinities 
within the Sterculiaceae, particularly for Theobroma and Herrania (Cuatrecasas, 1964, 
Contrib. U.S. Nat. Mus. 35:379-614), it would not be unexpected to discover in nature 
that herbivorous insects associated with one or the other genus successfully feed on the 
other genus as well. After all, there is at least one documented example of a genus of 
notodontid moth successfully exploiting both bombacaceous and sterculiaceous larval 
food plants (Young, J. Lepid. Soc. 37: 182-186), and these two Neotropical tree families 
are very closely related as well (Cronquist, 1981, An Integrated System of Flowering 
Plants, Columbia Univ. Press, New York). 

On 26 February 1988, I discovered two clusters of caterpillars on a “sapling” of H. 
albiflora (about 1.5 m tall and D.B.H. = 2.0 cm) planted along with a second individual 
of this species in a Theobroma and Herrania “garden” (n = approx. 30 trees for total of 
5 species) at “Finca Experimental La Lola,” near Siquirres (10°06’N, 83°30’W), Limon 
Province. The locality is within lowland tropical rain forest characteristic of the Atlantic 
watershed of Costa Rica. The locality experiences a short and irregular dry season be- 
tween December and March each year, although there are seldom days with no rainfall 
at all. 

When discovered one group contained 15 larvae all aggregated on the ventral side of 
two adjacent leaflets, and the second group had 10 larvae on a separate leaf. All larvae 
appeared to be 21-26 mm in body length, and based upon subsequent rearing data, were 
probably third or fourth instars. An eventual determination of the species resulted from 
rearing a sample consisting of one of the two groups, the second group being left undis- 
turbed on the food plant. Once collected and confined to a large clear-plastic bag kept 
tightly shut, the larvae were transferred to T. cacao leaves, in order to determine ac- 
ceptability of this species for successful development. The second group, left on the food 
plant, served as a control on this study. What initially led to the discovery of the larvae 
was the fact that the treelet was heavily defoliated, with more than half of the large, 
stellate-type leaves either completely missing (but not fallen off) or with only midribs 
remaining (Fig. 1). 

Caterpillars remain aggregated on leaves of both H. albiflora and T. cacao, although 
groups fragment into smaller clusters in the fifth instar (Fig. 1). Characteristically, in- 
dividual caterpillars feed from the already eaten edge of leaves and also rest in these 
positions when not feeding (Fig. 1). The leaves of Herrania are usually blotched in shades 
of light green and brown, a color combination that is matched by the mottled colors of 
the caterpillars (Fig. 1). Caterpillars feed and rest from loose webbings of silk spun over 
the leaves and lengthy petioles of Herrania leaves. 

A single mature leaf of H. albiflora consists of usually five leaflets, arranged in a stellate 
fashion. The “crown” of the treelet consists of a whorl of leaves in the absence of 
branches, and the very long petioles give the appearance of branches (Fig. 1). At the 
time of discovery, the treelet had a total of nine leaves, of which two were completely 
defoliated and the remaining ones with one to four leaflets missing on each (although 
five of the nine each had three leaflets missing). Thus the defoliation of the treelet by S. 
rustica was very advanced by this time. Absent were any new flushes of young leaves; 
no flowers and fruits were present. By 12 March (two weeks after initial discovery) only 
one of the original aggregate of ten caterpillars was left, and this individual was 45 mm 
long. Based upon the simultaneous rearing on T. cacao, this caterpillar (Fig. 1) was 
judged to be in the final instar. Presumably the others had matured and left the food 
plant for pupation by this time. The adjacent (about 3 m away) H. albiflora had no 
caterpillars, nor did it have any signs of the defoliation characteristic of this notodontid. 
The second tree, however, was in advanced stages of defoliation by an unidentified 
species of leaf-cutter ant, Atta sp. (Hymenoptera: Formicidae). Caterpillars kept on T. 
cacao thrived, many of these eventually pupating by 23 March while the sample was 
being hand-carried on an aircraft between Costa Rica and Nicaragua. Pupae are chest- 
nut-brown and range in body length from 18 to 20 mm. They eclose in 12-14 days, 
although the length of the pupal stage may vary greatly with temperature and other 


VOLUME 38, NUMBER 3 247 





Fic. 1. Caterpillar stage of Schizura rustica and associated defoliation activity. Be- 
ginning in upper left and clockwise: defoliated Herrania albiflora; three fourth instar 
caterpillars perched on partly devoured mature leaf; fifth instar caterpillar feeding and 
resting on partly devoured mature leaf (bottom two photographs). 


factors associated with husbandry. All adults eventually reared had been fed T. cacao 
following the time of the discovery. On H. albiflora in nature caterpillars were seen 
feeding during daylight hours, with intermittent periods of non-feeding. As typical for 
various notodontids, including temperate-zone Schizura (e.g., Packard, 1898, J. New York 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Entomol. Soc. 1:22-78; Forbes, 1948, Mem. 274, Cornell Univ. Agricult. Expt. Sta.), the 
caterpillars of S. rustica are exceedingly cryptic in coloration and in the habit of posi- 
tioning themselves along the damaged edge of a leaf on which they are feeding. 

What follows here is a general macro-description of the final instar caterpillar. Head 
capsule 4.5 mm high by 4.0 mm at widest lateral area; dull light brown background with 
heavy speckling of tiny dark brown spots laterally. Frontally with double band of vertical 
dark brown lines angling towards medial line about % down head capsule. Three pairs 
of frontal-to-lateral black setae about % down head capsule and a series of three pairs of 
clustered setae at base of head capsule in lateral areas. Double dark brown bands of 
frontal area forming a rough hourglass pattern when viewed frontally. Mandibles glossy 
brown and entire head capsule thrust forward basally. Light brown areas of head 
capsule speckled with very tiny brownish flecks, much lighter than those of dark brown 
areas. First thoracic segment forms a conspicuous “neck” ring with vivid yellow spot 
dorsomedially bordered laterally in very dark brown, almost black, diffuse bands. These 
bands give way more laterally to light brown. The dark brown bands bordering yellow 
spot each contain a small cream-colored spot. Below spiracle the segment is expanded 
posteriorly into wedge-shaped yellow flap. This segment with five pairs of black setae: 
one pair arising from dorsal dark brown bands; two pairs in lateral light brown area; two 
pairs arising from dorsal edge of spiracles. Legs light brown. Second and third thoracic 
segments light green with a prominent dorsomedial stripe that tapers near the end of 
the abdomen. This stripe is really a composite of a central thick yellow band bordered 
laterally with a mottled gray and brown area which is edged in a thin dark line of brown 
on either side. Green areas of these and other segments bearing tiny flecks of purple. 
Second thoracic segment with five setae when viewed laterally. Third segment begins a 
conspicuous rise in the dorsal area, forming a prominence that fuses with similar config- 
uration of first abdominal segment. Spiracles absent on second and third thoracic seg- 
ments. Legs on both segments also light brown. All abdominal segments various shades 
of brown. First abdominal segment light brown and mottled with a lacework of small 
dark brown lines; dorsally with a medial prominent biforked chitonous glossy peak. This 
prominence is reddish brown with one stout black seta oriented upwards on each. Spi- 
racular opening at the anterior border of this segment; ventrally all abdominal segments 
light green; two pairs of black setae readily visible laterally on first abdominal segment. 
The brownish lacework of abdominal segments 2—4 form a “saddle-like” configuration 
laterally; spiracles of these segments more centrally located on each side. Prolegs present 
on abdominal segments 3, 4, 5, and 6, and this entire region ventrally appears arched 
up; prolegs orange with reddish streaks. Three pairs of black setae on most abdominal 
segments, and most of these often arising from reddish bulbous basal structures. Abdom- 
inal background color becomes more blackish gray beginning with the third abdominal 
segment, developing into a broad dorsal band on the following three segments. The fifth 
abdominal segment bears a smaller dorsomedial prominence and continues on segments 
5 and 6, forming a second “peak” along the body axis. Dorsally the posterior half of the 
sixth segment is white, branching out into two bands on the seventh. The eighth segment 
with a very small dorsal prominence, and all three prominences biforked (as described 
above for the first “peak’’). Abdominal segment 8 with a lacework of dark brown lines 
as seen in first two abdominal segments, but now more reticulate over a light, reddish 
brown background. Spiracles ending on the eighth segment. The white dorsal area of 
the seventh segment, which branches laterally, abruptly ends with the eighth segment, 
and the dorsal area is colored with a blackish gray medial band again. This band contin- 
ues on the ninth and tapers into the anal clasper. The tenth segment is orange-brown 
and bears two pairs of black setae laterally, while the ninth segment has three pairs. Anal 
clasper orange with reddish terminal area. The caterpillar grows to about 45 mm prior 
to pupation. The overall appearance of the caterpillar, and its habit of resting on de- 
voured edges of leaves (Fig. 1), suggest a general strategy of crypsis, a trait presumably 
shared with earlier instars. 

Although some temperate-zone notodontids feed almost exclusively on young, soft 
tissue leaves of the larval food plants, others selectively feed on mature leaves (e.g., 
McFarland, 1979, J. Lepid. Soc. 33: Supplement). Aggregated feeding habits in the 


VOLUME 38, NUMBER 3 249 


caterpillars of some Lepidoptera, which often result in severe defoliation, are considered 
to be adaptations to food plant resources having very patchy distributions and therefore 
present in very limited supply (e.g., Tsubaki & Shiotsu, 1982, Oecologia 55:12-20; Fitz- 
gerald & Peterson, 1983, Anim. Behav. 31:417-423). At least one temperate-zone species 
of Schizura is polyphagous (Ferguson, 1975, U.S.D.A. Tech. Bull. No. 1521) and the 
possibility of such a habit being shared with S. rustica in Costa Rica cannot be ruled 
out. Both monophagous and polyphagous notodontids are known from the Neotropical 
Region (Seitz, 1907, Macrolepidoptera of the World, Stuttgart, A. Kernan). And, aside 
from the recent report of the notodontid Lirimiris meridionalis (Schaus), there are no 
other published accounts of Neotropical notodontids being associated with Sterculiaceae, 
and the present reports add a second genus and species to our knowledge of such asso- 
ciations. Given the close evolutionary affinities of Theobroma and Herrania (Cuatrecasas, 
op. cit.), the observed interchangeability of leaves from both trees to later instars of S. 
rustica is not a surprising or unexpected finding. Yet, in nature, other factors associated 
with the trees may select for egg-placement by this notodontid to be primarily a response 
to Herrania, a genus whose member species have leaf configurations and general tree 
profiles quite different from various Theobroma, including T. cacao. Noteworthy in this 
context is the fact that S. rustica was found only on H. albiflora, in spite of the fact that 
a handful of Herrania trees was surrounded by thousands of T. cacao (“cocoa’’) trees in 
a plantation setting. 

This research is a by-product of a research grant from The American Cocoa Research 
Institute. I thank Dr. Gustavo Enriquez and don Alfredo Paredes for local assistance at 
La Lola. The moth was identified by Dr. R. W. Poole of the U.S.D.A. Systematic Ento- 
mology Laboratory, and with the cooperation of Dr. Lloyd Knutson. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 249-251 


FOOD-PLANTS OF THE PIERIDAE 


After studying the correspondence between Messrs. Philip James DeVries and Allen 
M. Young (1982, J. Lepid. Soc. 36(3):299-232), it occurs to me that a general look at the 
food-plant preferences of the Pieridae, taken region by region, may be of interest. 


AFRICA 


Pseudopontiinae—No information. 
Coliadini 
Catopsilia—Cassia (Caesalpiniaceae), Sesbania (Papilionaceae), a somewhat dubious 
record of Gossypium (Malvaceae). 
Colias—Cassia (Caesalpiniaceae), Medicago, Phaseolus, Sesbania (Papilionaceae), Ox- 
alis (Oxalidaceae), a somewhat doubtful record of Ricinus (Euphorbiaceae). 
Eurema—Cassia (Caesalpiniaceae), Hypericum (Hypericaceae), Acacia, Albizzia, En- 
tada, Parkia, Dichrostachys (Mimosaceae), Aeschynome, Lespedeza, Sesbania (Pa- 
pilionaceae). 
Euchloini 
Pinacopteryx—Boscia, Cadaba, Capparis, Maerua (Capparidaceae). 
Euchloe—Barbarea, Iberis, Sisymbrium, etc. (Cruciferae). 
Pierinae 
Nepheronia—Ritcheia (Capparidaceae), Hippocratea (Hippocrataceae), Cassiporrea 
(Rhizophoraceae), Azima, Salvadora (Salvadoraceae). 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Eronia—Capparis (Capparidaceae), Salvadora (Salvadoraceae). 

Colotis—Boscia, Cadaba, Capparis, Maerua, Ritchiea (Capparidaceae) (some species 
eating both groups, some one only). A very dubious record of “dwarf bamboo in 
captivity.” 

Calopieris—No records. 

Gideona—No records. 

Belenois—Rhus (Anacardiaceae) (one species only), Boscia, Capparis, Cleome, Mae- 
rua, Ritchiea (Capparidaceae), Brassica (Cruciferae), Salvadora (Salvadoraceae), a 
very doubtful record of Solanum (Solanaceae). 

Pieris—Brassica (Cruciferae). 

Pontia—Alyssum, Brassica, Erucastrum, Lepideum, Sisymbrium (Cruciferae), Cay- 
lusia, Ochradenum, Reseda (Resedaceae), a dubious record of Solanum (Solanaceae). 

Dixeia—Capparis (Capparidaceae). 

Appias—Boscia, Capparis, Maerua, Ritchiea (Capparidaceae), Drypetes, Phyllanthus 
(Euphorbiaceae). 

Mylothris—Loranthus, Viscum (Loranthaceae), Osyris (Santalaceae) (one species along 
with Loranthus), Hevea (Euphorbiaceae), Theobromum (Sterculiaceae) (one species 
along with Loranthus), one species completely anomalous on Polygonum (Polygona- 
ceae). 

Leptosia—Capparis (Capparidaceae). 


ASIA (mainly India) and AUSTRALIA 


Coliadini 

Catopsilia—Bauhinia, Cassia (Caesalpiniaceae), Butea (Papilionaceae). 

Dercas—No records. 

Gonepteryx—Rhamnus (Rhamnaceae), Vaccinium (Ericaceae) (rarely). 

Gandaca—No records. 

Eurema—Caesalpinia, Cassia, Delonix, Wagatea (Caesalpiniaceae), Indigofera, Ses- 
bania (Papilionaceae). Also in Australia: Breynia, Phyllanthus (Euphorbiaceae), Al- 
bizzia, Leucaena, Pithecolobium (Mimosaceae). 

Colias—Astragalus, Oxytropis, Parochetus, Trifolium (Papilionaceae). 

Euchloeini 

Euchloe—Cruciferae spp. 

Pierini 

Leptosia—Capparis, Crataeva (Capparidaceae). 

Aporia—Berberis (Berberidaceae), Prunaceae spp., Rubiaceae spp. 

Delias—Loranthus (Loranthaceae), Nauclea (Rubiaceae), Averrhoa (Geraniaceae) (both 
the latter with the remark that the food-plant is more likely to be Loranthus growing 
thereon). 

Cepora—Capparis (Capparidaceae). 

Prioneris—Capparis (Capparidaceae). 

Anapheis—Capparis (Capparidaceae). 

Appias—Capparis, Crataeva (Capparidaceae), Hemicyclia (Euphorbiaceae). 

Pontia—Reseda (Resedaceae), Sinapis, Sisymbrium, Turritia (Cruciferae). 

Ixias—Capparis (Capparidaceae). 

Colotis—Cadaba, Capparis, Maerua (Capparidaceae), Azima, Salvadora (Salvadora- 
ceae). 

Hebomoia—Capparis, Crataeva (Capparidaceae). 

Valeria—Capparis (Capperidaceae). 


EUROPE 


Coliadini 
Catopsilia—Cassia (Caesalpiniaceae). 
Colias—Cistus (Cistaceae), Vaccinium (Ericaceae), Astragalus, Coronilla, Medicago, 
Trifolium, Vicia (Papilionaceae). 
Gonepteryx—Rhamnus (Rhamnaceae). 


VOLUME 38, NUMBER 3 Zl 


Euchloeini 
Euchloe—Barbarea, Biscutella, Ineris, Sisymbrium (Cruciferae). 
Anthocharis—Biscutella, Cardamines, Sisymbrium, etc. (Cruciferae). 
Pierini 
Aporia—Crataegus, Prunus, Spiraea (Rosaceae). 
Pieris—Aethionema, Alyssum, Brassica, Iberis, Sinapis, Sisymbrium (Cruciferae), 
Tropaeolum (Geraniaceae), Reseda (Resedaceae). 
Colotis—Capparis (Capparidaceae). 
Zegris—Sinapis (Cruciferae). 
Leptidea—Cracca, Lathyrus, Lotus, Viccia (Papilionaceae). 


> ce 


NORTH AMERICA (after Ehrlich & Ehrlich’s 


Coliadini 
Nathalis—Stellaria (Caryophyllaceae), Bidens, Dyssodia, Tagetes (Compositae), Ero- 
dium (Geraniaceae), Helenium (??). 
Colias—Vaccinium (Ericaceae), Amorpha, Astragalus, Hedysarum, Medicago, Paro- 
sela (Papilionaceae), Salix (Salicaceae). 
Kricogonia—No records. 
Eurema—Cassia (Caesalpiniaceae), perhaps Astragalus (Papilionaceae) and others. 
Phoebis—Cassia (Caesalpiniaceae). 
Euchloeini 
Anthocharis—Arabis, Barbarea, Cardamines, Sisymbrium (Cruciferae). 
Euchloe—Arabis, Sisymbrium, etc. (Cruciferae). 
Pierini 
Pieris—Dentaria, Isomeria, Stanleya, other Cruciferae and Capparidaceae. 
Ascia—Brassica, Cleome, Polanisia, other Cruciferae and Capparidaceae. 
Neophasia—Pinus (Coniferae). 


How to Know the Butterflies’) 


Unfortunately, I have no records for South America. 

Looking at the foregoing lists as a whole, a fairly coherent pattern emerges. The 
Coliadini are almost entirely confined to the leguminous subfamilies Papilionaceae and 
Caesalpiniaceae, with Gonepteryx confined to the Rhamnaceae. The other pierine tribes 
show a decided preference for plants containing mustard oil glucosides, i.e., Cruciferae, 
Capparidaceae and Salvadoraceae but with a few divergent groups or species; for ex- 
ample, Delias and Mylothris feeding mainly on Loranthaceae and Aporia on Rosaceae, 
Rubiaceae and Berberidaceae, among others. I am unable to trace any record for Lau- 
raceae apart from Mr. Young’s, and, although that does not completely preclude the 
family as a pierine food-plant, it makes it less likely. 


D. G. SEVASTOPULO, F.R.E.S., P.O. Box 95617, Mombasa (Nyali), Kenya. 


Journal of the Lepidopterists’ Society 
$8(3), 1984, 251-252 


ANTHOCHARIS LANCEOLATA (PIERIDAE) FEEDING ON A RARE 
ENDEMIC STREPTANTHUS SPECIES (CRUCIFERAE) 


Anthocharis lanceolata Lucas is recorded on several species of Arabis (Cruciferae) in 
various parts of its range. On 14 July 1983 it was found infesting the rare endemic 
Streptanthus howellii Wats. about 10 km southwest of O’Brien, Josephine County, Or- 
egon. The plants are on and adjacent to a disturbed roadside and power-line cut on a 
serpentine substrate and are confined to otherwise bare or nearly bare soil. Including 


252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


small vegetative individuals, there are at least 100 and possibly 200 plants in the area, 
making this one of the largest populations of S. howellii known. 

Streptanthus howellii is confined to dry, brushy serpentine exposures between 650 
and about 1000 m in the Siskiyou Mountains of Josephine and Curry counties, Oregon 
and Siskiyou and Del Norte counties, California. It is apparently a short-lived perennial 
or, perhaps, biennial. It is a candidate species for Federal listing under the Endangered 
Species Act, Category 2, USFWS (Federal Register 45:82480-82569, Dec. 15, 1980). 
Further information and figures are available (R. J. Meinke, 1982, Threatened and En- 
dangered Vascular Plants of Oregon, An Illustrated Guide, USFWS, Office of Endangered 
Species, Region 1, Portland, Ore., pp. 314-315). 

At the O’Brien site most of the large plants (about 15% of the population) bore one or 
more larvae of A. lanceolata. The single largest individual seen, a much-branched spec- 
imen over 1 m tall, had seven larvae and two eggs. Many of the smaller plants bore one 
or two eggs, generally on buds or on the axis of the raceme, rarely on leaves. Larvae 
were feeding actively on buds, flowers, and green fruit, and most of the siliques on the 
large plants were more or less damaged. A few plants had the entire inflorescence/ 
infructescence destroyed. Oviposition appeared to be correlated with phenophase, such 
that the most advanced plants bore the largest larvae; plants mostly in flower bore 
younger larvae and a few eggs; and plants in bud bore either eggs or nothing. No 
lanceolata were flying, and the latest plants to mature appeared likely to escape predation 
altogether. About a third of the eggs observed were dead, but the cause was not deter- 
mined. Eight larvae were brought back for rearing; seven pupated and one produced an 
undetermined braconid parasitoid (Hymenoptera). 

Meinke (loc. cit.) speculates that S. howellii may have “a poor reproductive capacity.” 
If this is the case, seed predation by pierid larvae, perhaps not limited to A. lanceolata, 
may be an important factor in its biology. At O Brien up to 75% of the seed crop appeared 
to be threatened (possibly less if the plants were able to regenerate and reproduce after 
the pierid feeding season) in 1983. Some other Streptanthus species on serpentine soils 
have evolved butterfly egg-mimics as an adaptation to predation-avoidance (Shapiro, 
1981, Amer. Nat. 117:276-294), but S. howellii does not have them, and it is not known 
whether A. lanceolata engages in egg-load assessment (though its eggs are typical of 
species which do). If other pierid-crucifer systems are at all typical, we may expect the 
impact of predation on S. howellii to vary drastically from year to year, depending on 
how weather modifies the phenology of the plants and insects. 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 252-253 


HACKBERRY BUTTERFLIES: DENSE SWARMS INVOLVED IN A 
LITIGATION IN SOUTHERN LOUISIANA 
(NYMPHALIDAE: ASTEROCAMPA) 


Hackberry butterflies (Nymphalidae, genus Asterocampa) are common insects of the 
central United States, ranging from southern New England westward throughout the 
mid-central United States to the Gulf of Mexico. They are especially abundant in the 
southern states of Arkansas, Texas, Mississippi, Louisiana, Alabama and western Tennes- 
see. In this region there are three annual broods beginning about May and extending 
into July, with the greatest number of insects occurring in June and July (Holland, 1947, 
The Butterfly Book, Rev. Ed., Doubleday and Co., Garden City, NY, 424 pp.; Klots, 


VOLUME 38, NUMBER 3 253 


1951, A Field Guide to the Butterflies of North America, East of the Great Plains, 
Houghton Mifflin Co., Boston, MA, 349 pp.). 

In common with other nymphalids, hackberry butterflies have long been known to 
occur in great clusters and swarms and to migrate. Their presence in Louisiana has been 
well documented on several occasions (Lambremont, 1954, Tulane Stud. Zool. 1:127- 
164; Ross & Lambremont, 1963, J. Lepid. Soc. 7:148-158). The fact that they occur in 
enormous population densities in the mid-southern United States has been recorded as 
long ago as 1888. For example, in May of that year enormous numbers were noted in 
flight, and the banks of the St. Frances River in Arkansas were reported to be lined with 
Asterocampa celtis Boisduval and Leconte for a distance of over thirty miles (Webster, 
1888, Holtzgang, 1888, Insect Life 1:29 cited in Williams, 1930 below). In that same 
year it was reported that hackberry butterflies were swarming in great numbers over the 
southern United States and appeared to be migrating in a southerly direction (Williams, 
1930, The Migration of Butterflies, Biol. Monogr. and Manuals Nr. 9, Edinburgh, Oliver 
and Boyd, London, 4783 pp.). 

In the months of June and July of 1980 I noted a very large swarm of hackberry 
butterflies in East Baton Rouge Parish, Louisiana. A particularly dense cluster was ob- 
served over a period of about three weeks in the southern part of the city of Baton Rouge. 
Several thousand could be seen at any given time throughout any sunny day in my 
backyard. The insects were clustered on fig trees, feeding on the ripe fruit. Some indi- 
vidual figs often had five to ten butterflies resting and feeding. Many alighted on me 
while I spent many hours watching their behavior and abundance. Both A. celtis and A. 
clyton Boisduval and Leconte were identified, with A. celtis being far more abundant, 
comprising about 90% of the total swarm. The behavioral habit of hackberry butterflies 
alighting on people, even as they work, has been noted in the earlier literature during 
dense swarms of these insects (Williams, 1930, op. cit.). 

About a year later I was approached by a local attorney to identify an insect that 
factored in litigation between a homeowner and a painting contractor in West Baton 
Rouge Parish, Louisiana, directly across the Mississippi River from the City of Baton 
Rouge. The suit by the homeowner was to recover damages and cost of correcting poor 
workmanship in the exterior painting of a house. Large numbers of insects were stuck 
in the paint, and the painting workmen stated that they could not complete their work 
on the date promised because of large swarms of “moths” that were alighting on them 
and the freshly painted surfaces. 

The homeowner had taken numerous color photographs of the house including many 
close-up views of the insects themselves. Since several thousands of dollars were involved, 
I was asked to serve as an expert witness and was provided a photograph dated July 
1980. The photograph revealed intact male and female specimens and many detached 
wings, legs, antennae, and scales of Asterocampa celtis. From statements made at the 
trial, this species was swarming in numbers even greater than I had observed in the city. 
Their density and swarming and their behavior pattern of alighting on people as brought 
out in the hearings and well documented in the scientific literature, played a mitigating 
role for the defendants in this case. 


EDWARD N. LAMBREMONT, Nuclear Science Center, Louisiana State Univeristy, Ba- 
ton Rouge, Louisiana 70803-5820. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 254-256 


BOOK REVIEW 


PHENETICS AND ECOLOGY OF HYBRIDIZATION IN BUCKEYE BUTTERFLIES (LEPIDOPTERA: 
NYMPHALIDAE). John E. Hafernik, Jr., University of California Publications in Entomol- 
ogy, Volume 96. 118 pp., 35 line drawings, 15 halftones, February 1983, $16.50, ISBN 
0-520-09649-5. 


This work analyzes the ecological and phenetic ordinant relationships of Junonia in 
North and Central America, exclusive of the Caribbean. Hafernik assesses competition 
in the field between J. coenia, J. nigrosuffusa, and J. zonalis and infers their genetic 
relationships. The text has 42 pages divided into five sections: Intraspecific and Interspe- 
cific Crosses; Courtship Behavior; Population Size, Vagility and Dispersion of South Texas 
Junonia; Larval Resource Partitioning; and Phenetics. 

Hafernik notes that while electrophoretic assays of enzyme variability allow quanti- 
tative estimates of genetic differentiation between taxa, he prefers hybridization studies, 
because they illuminate hybrid fitness via egg fertilities, embryo viabilities, skews of sex 
ratios and progeny mortality. He investigates these relationships within Junonia by cross- 
ing F, and F, hybrids, and backcrossing among J. coenia from California and Texas, J. 
nigrosuffusa from Texas, and J. zonalis from Guatemala. Hybrid matings were obtained 
by substituting a different female in the middle of a natural courtship. Data for egg 
fertility, egg viability and percent of hatch were not statistically analyzed. Data on sex 
ratios were analyzed using chi-square to compare both individual broods and pooled 
values of like broods with an expected 1:1 ratio. Noncontrolled rearing environments 
precluded quantitative comparisons of generation times, but these data, as well as mor- 
tality estimates, emergence synchronies of the sexes, and incidences of aberrations were 
compared qualitatively. 

Hafernik’s hybridization data suggest that North and Central American Junonia are 
one polytypic species rather than a circle of races, since interpopulation genetic com- 
patibility is high regardless of geographic distances. These findings are in contrast with 
other studies of papilionoids; Hafernik reviews many similar studies. 

Hafernik states that Junonia lack complicated courtship rituals. Males rest on a bare 
spot on the ground and pursue suspect females pugnaciously. Visual stimuli, especially 
background color of the dorsal wing surfaces, appear to trigger male responses. Hafernik 
tested male response to females using various wing marking and obscuring techniques 
and also tested models and wing transplant females. Color differences limiting coenia 
and nigrosuffusa courtship interactions are considered unrelated to either thermoregu- 
lation or crypsis. Aposematicity was not tested (Junonia and Euphydryas share similar 
hosts in the Scrophulariaceae with iridoid glycosides). There is no evidence of pheromone 
involvement in Junonia, although no experimentation was carried out in this vein. 

Hafernik’s estimates of population dynamics and vagility for Junonia are based on 
populations at Brazos Island, Texas. Mark/recapture studies followed Ehrlich and Da- 
vidson (1960, J. Lepid. Soc. 14:227-230), with vagility analyzed between two sectors over 
distances of ca. 2 km using Scott’s (1972, Ph.D. thesis, U. of Calif., Berkeley) technique. 
Jolly’s (1965, Biometrika 52:225-247) method was used for estimates of population. Dis- 
persion was analyzed using the variance mean ratio and Morisitas indices (Southwood, 
1966, Ecological Methods, Methuen, London). Hafernik concludes that coenia and ni- 
grosuffusa have similar vagility patterns with males markedly more aggregated due to 
mating and females more dispersive for host selection. The micro-distribution of females 
in the environment is different for these species, with nigrosuffusa using and spending 
more time at clumped host plants, while coenia spend more time in transit between 
unclumped hosts. Males showed similar highly contagious distributions but had little 
spatial overlap. Male coenia chose short vegetation for loitering, while nigrosuffusa chose 
taller vegetation, chiefly stands of sedges. Hafernik postulates that coenia males may 
have a competitive advantage in short vegetation, based upon sympatric interaction with 
male nigrosuffusa and upon the latter’s behavior and mating area choice, in Arizona 
under allopatric conditions. 


VOLUME 38, NUMBER 3 25a 


Studies on larval resource partitioning involved eight localities in Texas over three 
years and one location in Arizona. While larvae of coenia and nigrosuffusa could not be 
distinguished, reared adults were identified using Discriminant Function Analysis (DFA) 
(see below). Host palatability was tested by presentation of hosts to allopatric populations 
of larvae of coenia, nigrosuffusa and zonalis. Adult female oviposition preferences in 
cages were also noted. Hafernik’s data indicate considerable host overlap in south Texas 
with coenia chiefly using Agalinis maritima, and nigrosuffusa using Stemodia tomen- 
tosa in the presence of coenia but also A. maritima in allopatry. J. nigrosuffusa shows 
better larval development on the latter, whereas, coenia is limited by the leaf pubescence 
of Stemodia. Hafernik postulates that the perennial and annual habits of Stemodia and 
Agalinis, respectively, may account for falling numbers of adult coenia in winter, while 
adult nigrosuffusa populations remain high. He speculates on the implications of this 
regarding hybrid introgression. J. zonalis from Guatemala showed more restricted host 
preferences than coenia or nigrosuffusa, and ovipositing females rejected the favored 
hosts of the latter two. 

In the Phenetics section, Discriminant Function Analysis (DFA) and Principal Com- 
ponent Analysis (PCA) were used to demonstrate the relationships between known pa- 
rental and hybrid reference groups and between reference groups versus unknowns from 
Mexico and Central America. Hafernik chose 25 wing characters (7 continuous, 17 coded 
[discrete]) for his analyses. While he states that the coded characters violate the para- 
metric assumptions of the DFA’s and PCA’s, he notes: (1) a statement by Blacklith and 
Reyment (1971, Multivariate Morphometrics, Academic Press, New York) that DFA and 
PCA are “robust” enough to handle [minor] violations of normality, and (2) that the 
derived DFA classifications of F, hybrids were empirically correct (his figure 18). Tra- 
ditionally, however, inclusions of nonparametric data in such analyses account for less 
than 20 percent of characters. In Hafernik’s work, they equal 68 percent and dominate 
the vectors. Thus, the discriminators chosen by the DFA are specifically antagonistic to 
the assumptions of the analyses. This bothers me, though the DFA scatter-plots are 
undoubtedly plausible. Interestingly, Hafernik (p. 39) states that principal component 1 
of the PCA (“unlike DF1I*’) shows moderate to high loadings for all coded characters; 
this demonstrates the sensitivity of the analyses to variation differences between the 
continuous versus coded characters used. The use of only continuous characters for the 
DFA would have avoided this philosophical conflict, as would have the use of Principal 
Coordinate Analysis or Nonmetric Multi-dimensional Scaling instead of PCA, if coded 
characters were retained. 

Hafernik infers genetic relationships based upon DFA and PCA results, citing empir- 
ical evidence for this conclusion via multivariate analysis examples of Rohwer (1972, 
Syst. Zool. 21:313-338), Rohwer and Kilgore (1973, Syst. Zool. 22:157-165), Jackson 
(1973, Evolution 27:58-68) and Thaeler (1968, Evolution 22:543-555). These authors, 
however, followed traditional constraints in using nonparametric data in DFA and PCA. 
Hafernik’s apparent major deviation contracts the logical consistency employed by those 
he cites. Not that I doubt the probability of biological (genetic) correctness of Hafernik’s 
results (it is hard to argue against “proven” empiricism), but rather I find the results an 
analytical curiosity and testament to the “robustness’’ of multivariate analyses. One could 
argue, however, that in the transparent guise of “robustness” of statistical methods, the 
philosophy that “the end justifies the means” is a bit too visible. 

In essence, I found Hafernik’s work excellent biology and very interesting reading. 
The text is lucid and Hafernik reviews his subjects well during discussion. Seldom is this 
much information produced on a subject such as hybridization unless team efforts are 
involved. The work leaves a hunger for the answers to those inevitable questions one can 
ask only when well into research. I only hope Hafernik or another population ecologist 
will explore the other side of this coin—the electrophoretics. 

On the negative side, some of the graphics could have been improved, especially 
labeling on histograms and scattergrams, and as stated above, parametric variables should 
have been used in the phenetic analyses. 

Interestingly, the text of this volume is typeset, unlike some previous University of 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


California Publications in Entomology serials, which were typewritten. I hope this un- 
predictable luck continues, since appearance alone does have implications for the quality 
of any series (are you listening U.C. Press?). 

At $16.50, Hafernik’s work is well worth its price and will be necessary for any “lep’er” 
who claims to be a biologist or biologist who researches leps. 


J. T. SORENSEN, Insect Taxonomy Laboratory, California Department of Food and 
Agriculture, Sacramento, California 95814. 


Journal of the Lepidopterists’ Society 
38(3), 1984, 257-258 


OBITUARY 


HAMILTON ALDEN TYLER (1917-1983) 


Hamilton A. Tyler, a versatile scholar, died at his home in Healdsburg, California, on 
14 December 1983. He was born in Fresno on 20 October 1917, the son of Hazel Tyler 
and John G. Tyler, the latter having been a well-known ornithologist of the Fresno area. 

Tyler was known in recent years for his studies of the Papilionidae of the New World: 
“The Swallowtail Butterflies of North America” (1975). At the time of his death he was 
writing, with Paul Spade, an article on Colima swallowtails. This manuscript is being 
prepared for publication by Michael Parsons under the title of “Notes on the Biology of 
Seven Troidine Swallowtail Butterflies (Lepidoptera: Papilionidae) from the State of 
Colima, Mexico.” 

Hamilton Tyler’s first entomological studies were on the Tenebrionidae, especially the 
beetles of the genus Eleodes, of which he had a large collection. As a young man, after 
studies at the University of California, Berkeley, and an episode as a soldier in the Spanish 
Civil War, he returned to the University as an English major and published papers on 
the poetry of Milton and Donne. 

In 1955 he became interested in the Southwest and began a study that resulted in 
three books on Pueblo myths that were published by the University of Oklahoma in their 
Civilization of the American Indian series. 

Tyler’s interest in plants led to several published articles on geraniums, two books on 
gardening and the editorship of a quarterly international publication called “The Pel- 
argonium Breeder.” 

He was also interested in birds, which led to the publication of Owls by Day and 
Night with Don Phillips in 1978. He and Phillips had also prepared, but had not yet 
published, a detailed manuscript on the owls of the Southwest and Mexico. 

Following his death, his collection of swallowtails and reference materials pertaining 
was given to the University of California at Davis. Of this collection, Dr. Arthur M. 
Shapiro writes: “Hamilton Tyler was an extremely active amateur lepidopterist who did 
much to foster research on the biology and systematics of his favorite group, the Swal- 
lowtails (Papilionidae). His collection, obtained by collaboration, purchase, and exchange, 
represents one of the largest accumulations of this family in private hands in North 
America. It is most notable for containing specimens with full data of many very rare 
and prized entities.” 

Dr. Shapiro also writes: “Basically, acquisition of the Tyler collection allows scholars 
and researchers in the region and in general to pursue evolutionary and phylogenetic 
studies of this important group without having to go to the British Museum, U.S. National 
Museum, or Allyn Museum (Sarasota, Florida). Mr. Tyler was working on a book of the 
Papilionidae of the Americas at the time of his death; most of the documentation is in 
the collection. He knew the group well enough to be a connoisseur and to obtain the 
‘key’ species for his work, which are likely to be critical for future workers as well.” 

The variety of publications by Tyler is best explained in the following statement by 
him: “Almost all of my books have been on topics of my own choosing and were thereby 
expressions of deep personal interest rather than material written to fit a publisher's 
demands. By good fortune on this chancy route, all completed titles have eventually 
reached publication despite some tortuous turns and delays. My interests have been 
diverse, as the varied titles indicate, and I have dug in some unusual corners but I think 
there is an overall logic which relates the works. My primary interest is in examining 
the different views men may take of the world about them, especially as these concern 
birds, mammals, insects, and plants, or relationships in the realm of nature. These ap- 
proaches seem to fall into two great and familiar divisions which are often held to be 
mutually exclusive—but they are not so for me. On the one side there is the scientific 
attitude which distinguishes the species of bird, mammal, insect or plant, by describing 
its form, life history, and habitat. On the other side there are views, such as those held 
by the Pueblo Indians, which are more poetic. Collectively these can be called the 


258 





JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mythologic approach to nature and on this side stand religions, at least until they become 

highly sophisticated. For me, both the mystic-poetic and the scientific outlooks illuminate 

man’s place in nature and the cosmos and are thus equally valid. Indeed, the two views 

complement each other. Both aesthetics and science are needed to establish man’s place, 

for they are the two hemispheres of the one world we all inhabit.” 
The following is a chronological listing of his publications: 


PUBLICATION 


Mr. Eliot and Mr. Milton 


Finnegan Epic 
Pueblo Gods and Myths 


Organic Gardening Without Poisons 

French Type of Tetraploid “Geraniums”’ in 
The Geranium Gazette Yearbook 

John Tyler (with John R. Arnold) 


Gourmet Gardening 


Pueblo Animals and Myths 
The Swallowtail Butterflies of North America 
Owls by Day and Night (with Don Phillips) 


Pueblo Birds and Myths 


PUBLISHER 


Circle, Berkeley 
Circle, Berkeley 
University of Oklahoma 
Van Nostrand Reinholt 


Auk, vol. 88:228-229 
Van Nostrand Reinholt 
University of Oklahoma 
Naturegraph 
Naturegraph 

University of Oklahoma 


JOHN R. ARNOLD, 199 Calistoga Rd., Santa Rosa, California 95405. 


DATE 


1944 
1946 
1964 
1970 
1971 


1971 
1972 
1975 
1975 
1978 
1979 


Journal of the Lepidopterists’ Society 
38(3), 1984, 259-260 


OBITUARY 


IN MEMORIAM 
RICHARD FABIAN TOWNSEND (1939-1983) 


Dr. Richard Fabian Townsend died 12 September 1983 in Anderson, California. He 
was born 8 November 1939 in Los Angeles, California and lived in the state all his life. 
In 1962 he married Diane Moseman of Canoga Park, California. He received an A.A. 
degree in pre-pharmacy from Los Angeles Valley College, then his Pharm. D. from the 
University of Southern California. He worked as a pharmacist for Thrifty Drugs in 
Redding, then was a pharmacy consultant for the state. His last employment was as a 
biological technician for the U.S. Forest Service, Redding. 

I first met Dick in 1955 at the Lorquin Entomological Society, Los Angeles County 
Museum of Natural History, where so many budding professional and amateur ento- 
mologists have received enlightenment. Dick and I were especially encouraged by Mr. 
Lloyd Martin and Dr. Fred Truxal of that institution. We took our first extended field 
trip together in June 1955: eight days (without a bath!), sleeping in a pup tent, while 
chasing such “wonders” as Colias eurydice, Papilio bairdii bairdii and Speyeria coronis 
semiramis near Barton Flats in the San Bernardino Mountains of southern California. I 
will never forget our efforts to prevent ground squirrel maraudings into our food supply, 





Dr. Richard Fabian Townsend 


260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


nor the expletive-charged argument as to who was at fault for our failure to net the only 
Baird’s swallowtail seen! Later that summer, and again the following one, my mother 
sacrificed considerable time and effort to become “expedition leader” while Dick and I 
chased butterflies and beetles over much of Arizona. Highlights of those trips include 
Dick misplacing our only road map into the ice chest and his animated (to put it mildly) 
reaction to being stung on one of his private parts by an angry harvester ant! These are 
just some of the many memories we enjoyed over the years. 

During succeeding years, until about 1961, Dick and I collected together and in the 
company of others, mostly on short trips in southern California but also as far afield as 
southeastern Arizona. Shortly thereafter, Dick made two extensive collecting trips to 
Mexico; one by bus. By this time Dick’s interests tended more towards Coleoptera. He 
donated most, if not all, his earlier butterfly collection to the Universidad Nacional 
Aut6énoma de México, under the care of Dra. Leonila P. Vazquez G. 

Dick and I enjoyed a special camaraderie during those early years of our friendship, 
one which decreased only spatially as we pursued disparate education and careers. Dur- 
ing Dick’s years of practicing pharmacy his interest in entomology gave way largely to 
such pastimes as fishing, hunting and prospecting for old bottles and other artifacts. 
However, during his employment with the U.S. Forest Service his “first love” was rekin- 
dled and butterflies came to be his primary objective. We kept in touch over the years 
and, especially during the last two, were able to renew our collecting together. 

The Dick Townsend Collection will be donated to the Los Angeles County Museum 
of Natural History. It comprises a specialized collection of approximately 250 Lepidop- 
tera, many of which were reared, including some excellent material in the genus Papilio; 
and 2800 Coleoptera, almost entirely in the families Buprestidae, Cerambycidae and 
Scarabaeidae. 

Dick is survived by his lovely wife and daughters (Erin and Stacie), his mother, Erma 
Townsend and sister, Linda Hensley. He was my friend of longest standing. We will all 
miss him very much. So too will his fellow lepidopterists. 


RICHARD L. WESTCOTT, Oregon Department of Agriculture, Salem, Oregon 97310- 
0110. 


Date of Issue (Vol. 38, No. 3): 24 April 1985 


EDITORIAL STAFF OF THE JOURNAL 
THomas D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MAGDA R. Papp, Editorial Assistant 
DouGLas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 
NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name, with authors of 
zoological names. Insect measurements should be given in metric units; times should be 
given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where 
italics are intended. References to footnotes should be numbered consecutively, and the 
footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 196la, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Ady. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 28-30). 

Illustrations: All photographs and drawings should be mounted on stiff, white back- 
ing, arranged in the desired format, allowing (with particular regard to lettering) for 
reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches 
are not acceptable and should be reduced photographically to that size or smaller. The 
author’s name, figure numbers as cited in the text, and an indication of the article’s title 
should be printed on the back of each mounted plate. Figures, both line drawings and 
halftones (photographs), should be numbered consecutively in Arabic numerals. The term 
“plate” should not be employed. Figure legends must be typewritten, double-spaced, on 
a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, 
with a separate paragraph devoted to each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material should be kept to a minimum and 
must be typed on separate sheets, and placed following the main text, with the approx- 
imate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of 75¢ per line. A purchase order for reprints will accompany the 
proofs. 

Correspondence: Address all matters relating to the Journal to the editor. Short 
manuscripts such as new state records, current events, and notices should be sent to the 
editor of the News: June Preston, 832 Sunset Drive, Lawrence, Kansas 66044 U.S.A. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


SOD WEBWORM MOTHS (PYRALIDAE: CRAMBINAE) IN SOUTH DA- 
KOTA. B. McDaniel, G. Fauske ¢& R. D. Gustin 0. . 149 

PAPILIO EURYMEDON LUCAS, 1852: A SYNONYM OF PAPILIO 
ANTINOUS DONOVAN, 1805 (PAPILIONIDAE). Murray S. 


Upton oe ee 165 
HAMADRYAS IN THE UNITED STATES (NYMPHALIDAE). Dale W. 
Jenkins 2 ee nit 


A SEX PHEROMONE IN THE CALIFORNIA OAKWORM PHRYGANIDIA 
CALIFORNICA PACKARD (DIOPTIDAE). Michael E. Hoch- | 
berg dr W. Jan A. Volney 176 


POPULATION BIOLOGY OF THE GREAT PURPLE HAIRSTREAK, AT- 
LIDES HALESUS, IN TEXAS (LYCAENIDAE). Paul L. Whit- 
taker jo i i . 179 


FORAGING BEHAVIOR OF TAWNY EMPEROR CATERPILLARS 
(NYMPHALIDAE: ASTEROCAMPA CLYTON). Nancy E. 
Stamp 20 ee 186 


Host SHIFT OF ECPANTHERIA DEFLORATA (ARCTIIDAE) FROM AN’ 
ANGIOSPERM TO A LIVERWORT. Kevin C. Spencer, Larry 
R. Hoffman & David S. Seigler ____.___.__._ a 192 


ETHOLOGY OF DEFENSE IN THE APOSEMATIC CATERPILLAR PA- 
PILIO MACHAON SYRIACUS (PAPILIONIDAE). David L. 
Beans) 194 


THE EGG OF HOFMANNOPHILA PSEUDOSPRETELLA (OQECOPHORI- 
DAE): FINE STRUCTURE OF THE CHORION. Richard T. Ar- 
bogast, Richard Van Byrd, Georges Chauvin & Rudolph G. 
Strong) 200 ee 202 


THE DYNAMICS OF ADULT DANAUS PLEXIPPUS L. (DANAIDAE) 
WITHIN PATCHES OF ITS FOOD PLANT, ASCLEPIAS SPP. M. 


P. Zalucki ts R.L. Kitching Ee 209 
BUTTERFLIES OF Two NORTHWEST NEW MEXICO MOUN- 
TAINS. Richard Holland Ee 220 


GENERAL NOTES 
The larch casebearer, Coleophora laricella (Hiibner) (Coleophoridae), in 


western Washington. Sanford R. Leffler 0 235 
The Gesneriaceae and Bignoniaceae as food-plants of the Lepidoptera. D. 
G: Sebastopulo 235 
Notes on the natural history of Papilio victorinus Doubl. (Papilionidae) in 
northeastern Costa Rica... Allen M. Young 1 237 
“Edge effect” in oviposition behavior: a natural experiment with Euchloe 
ausonides (Pieridae). Arther M. Sharir occccceeceeccceeesnvncccsssnnceseeveseeseesnsseeseeesesee 242 
Epiblema luctuosana A. Blanchard, a homonym, is changed to Epiblema 
luctuosissima, new name. André Blarrchar ooiceeeecccecsncceseseecceeeenneesnsensseseerseee 245 
Schizura rustica (Schaus), a notodontid moth defoliating Herrania and Theo- 
broma species (Sterculiaceae) in Costa Rica. Allen M. Young ............. . 245 
Food-plants of the Pieridae. D. G. Sevastogrlo oi cecccecsnccccceesssncceeeeesssnnesesersuseneees 249 
Anthocharis lanceolata (Pieridae) feeding on a rare endemic Streptanthus 
species (Cruciferae). Arthur M. Shapiro 0 251 
Hackberry butterflies: dense swarms involved in a litigation in southern Lou- 
isiana (Nymphalidae: Asterocampa). Edward N. Lambremont 0c 252 
BOOK REVIRW sk a i 254 


OBITUABIES ois he a a een 257, 259 


2m hen) Ser aa 


Number 4 


1984 


Volume 38 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ 


SOCIETY 


THE LEPIDOPTERISTS’ SOCIETY 


by 
Publié par LA SOCIETE DES LEPIDOPTERISTES 


Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


Published quarterly 


Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


ove 
Ct he 





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Seah Ss 
‘ (| = 
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Cans he) 


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10 July 1985 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Don R. Davis, President LEE D. MILLER, 

ViToR O. BECKER, Vice President Immediate Past President 
JAVIER DE LA MAZza E., Vice President JULIAN P. DONAHUE, Secretary 
JOHN C. DOWNER, Vice President Eric H. METZLER, Treasurer 


Members at large: 


F. S. CHEW J. M. BURNS B. A. DRUMMOND 
G. J. HARJES F. W. PRESTON J. LANE 
E. H. METZLER N. E. STAMP R. K. ROBBINS 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229, U.S.A.; and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$25.00 (institutional subscriptions) and $18.00 (active member rate) by the Lepidopter- 
ists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Boule- 
vard, Los Angeles, CA 90007. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, 
1900 John St., Manhattan Beach, CA 90266. 


Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- 
temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- 
sion in the Sesiidae fascicle for the Moths of America North of Mexico. The dusky 
clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of 
willows and poplars. 


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JOURNAL OF 


Tue LeEpIpoPTERISTS’ SOCIETY 











Volume 38 1984 Number 4 





Journal of the Lepidopterists’ Society 
38(4), 1984, 261-267 


HYBRIDIZATION BETWEEN CALLOSAMIA AND 
HYALOPHORA (SATURNIIDAE) 


THOMAS W. CARR 
6626 Weckerly Drive, Whitehouse, Ohio 43571 


ABSTRACT. Several intergeneric crosses involving Callosamia and Hyalophora were 
attempted. Male and female F, adults were obtained from the cross C. angulifera 6 x 
H. cecropia 2. All stages were intermediate, exhibiting characteristics of both parent 
species. Other crosses, which did not produce adults, are discussed. 


Although Ferguson (1972) restored Callosamia and Hyalophora to 
full generic rank, he acknowledged that they were undoubtedly closely 
related. In spite of this apparent close relationship I know of no natural 
intergeneric hybrids. The only attempts to artificially induce hybrids 
are those mentioned by Peigler (1978) and Collins and Weast (1961). 
Efforts to obtain intergeneric hybrids might produce information which 
would help clarify the relationship between Callosamia and Hyaloph- 
ora. In this paper I describe my hybridization attempts and results. 
The discussion includes a comparison of these results with those of 
other hybridization studies invoiving these genera and Samia cynthia 
(Drury). 


METHODS AND MATERIALS 


In June 1979, using hand-pairing techniques described by Peigler 
(1977), a male Callosamia angulifera (Walker) from Boone Co., West 
Virginia was mated to a female Hyalophora cecropia (L.) from Lucas 
Co., Ohio. The moths were transferred to a foothold where they re- 
mained coupled for ca. three hours, after which the female oviposited 
freely in a paper sack. 

Three additional matings of the same combination were subsequent- 
ly obtained. Often, the movement of the very large females threatened 
to dislodge the males. This was prevented by placing the female on a 
flat surface and pinning paper strips over the folded wings. Pins were 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


also placed at strategic points alongside the female’s abdomen to fur- 
ther minimize movement. After copulation, females were placed in a 
common container where they oviposited freely. 

Resultant larvae were reared on tuliptree (Liriodendron tulipifera 
L.). Large cloth bags (sleeves) were placed over branches with the 
larvae confined within. 


RESULTS 


The first female, which was confined separately, deposited nearly 
200 ova, 33% of which hatched. Unfortunately, because the remaining 
three females were confined in the same container, variation in fertility 
could not be determined. Of the 600 ova deposited hatch was again 
33%. Upon eclosion I retained 150 of the larvae and gave the remain- 
der to Dana Gring, Toledo, Ohio. His results were similar to those 
described in this paper, but I have no specific data. 

Most larval losses occurred in the first and second instars. Disease 
did not appear to be a major factor in these losses. Unidentified pre- 
dacious stinkbugs (Hemiptera: Pentatomidae) pierced and killed larvae 
from outside the rearing sleeves. Later attacks were prevented by cov- 
ering the first sleeve with a second one. 

A total of 74 cocoons was obtained. Six females and 36 males emerged 
the following summer. The remaining cocoons contained either dead 
pupae or females that were unable to escape their cocoons. 

A brief description of the various stages follows: First and second 
instar larvae appeared structurally similar to H. cecropia. Color be- 
came lighter with age, eventually more closely resembling C. anguli- 
fera. The third and fourth instars appeared much more intermediate 
structurally and in overall color, basically resembling the mature larva. 

The fifth instar larva (Fig. 1) had the ground color blue-green. The 
first two pairs of thoracic scoli were deep red, bearing minute setae; 
the third pair was orange with yellow bases. The remaining dorsal scoli 
were lemon-yellow. The subdorsal and subspiracular tubercles ap- 
peared as raised points, varying in color from dark to light blue in 
different individuals. The yellow subspiracular stripe found on C. an- 
gulifera was absent. 

Pupae were intermediate in size. The brown color was very close to 
that of C. angulifera. Cocoons were also intermediate in size, averaging 
5.3 cm in length, with a double wall as in both parent species. Color 
was dark brown and uniform in all examples. Two larvae attached 
their cocoons to branches lengthwise as in H. cecropia. Two others 
spun weak leaf stem attachments; all remaining larvae spun unattached 
cocoons amongst leaves or in folds of rearing sleeves. 

The adult male (Figs. 2, 3) had antennae intermediate in size with 


VOLUME 38, NUMBER 4 263 





ws Fe r 2 a 8 Maes 


Fic. 1. Hybrid fifth instar larva from C. angulifera 6 x H. cecropia 8. 


the medium brown coloring like C. angulifera. The body was dull 
wine-red. The prothoracic collar was usually a poorly defined gray, 
and a few had some white shading. White segmental rings were present 
on the abdomen, as in H. cecropia. The ground color of the wings was 
dark brown overlaid with a wine-red cast. Grayish suffusion was lim- 
ited to the forewing costa. Antemedial lines were intermediate, more 
prominent than in C. angulifera, and sharply angled on the forewing, 
with an inner white shading, as in H. cecropia. Each discal spot had a 
prominent anterior tooth. The white postmedial line was shaded out- 
wardly with purplish pink, this color being more diffuse than the sharp- 
ly delineated red of H. cecropia. The overall coloring of the underside 
was red-brown inside of the postmedial line and had a pink shade 
outwardly; the overall aspect being reminiscent of C. angulifera. The 
underside of the hindwing had a wide white costa. 

A genitalia study (Fig. 5) of three males indicated a complete de- 
velopment of the aedeagus which failed to exhibit a distinguishable 
tendency toward either parent species. In two of the specimens the 
valvae could best be described as shapeless due to a lack of scleroti- 
zation, which may or may not be an artifact of preparation. The re- 
maining male possessed genitalia with an exaggerated development of 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 2-4. Adult hybrids from C. angulifera 6 x H. cecropia 2. 2, male (dorsal view, 
upper left); 3, male (ventral view, upper right); 4, female (dorsal view, lower). 


the median lobes of the valvae into long and pointed processes which 
are characteristic of C. angulifera. 

Two female forms were obtained. In the first form (Fig. 4) the 
ground color of the body and wings was bright reddish brown, dorsal 
thoracic collar gray, and white segmental rings of the abdomen not as 
prominent as in the male. The antemedial and postmedial lines were 
as described for the male; the anterior tooth of each discal spot was 
present but not as prominent as in the male; underside was similar to 
the male but lighter brown in color. The second form (one specimen) 
was considerably larger than the others, with a ground color of light 
brown with no reddish cast; the anterior tooth of each discal mark was 
barely present, resembling H. cecropia. The white abdominal rings 
and lateral chain-like ornamentation of the abdomen was barely dis- 
cernable; the pink shading of postmedial lines was very faint, nearly 
absent. Each female contained very few ova. 

Hybridization attempts involving other combinations of species within 
these genera produced no adults. The cross H. cecropia 6 x C. angu- 


VOLUME 38, NUMBER 4 265 





Fic. 5. Male genitalia of hybrid from C. angulifera 6 x H. cecropia &. 


lifera 2 produced two larvae which resembled pure C. angulifera. 
They were very weak and did not feed, expiring after a few hours. 
The cross H. cecropia 6 x C. promethea (Drury) 2 produced one larva 
which fed on common chokecherry (Prunus virginiana L.) for three 
days before expiring. Ova from the reciprocal cross did not hatch. 


DISCUSSION 


Hybrid males were vigorous and easily escaped their cocoons. As 
previously noted, the majority of females failed to emerge, apparently 
being too weak to do so. Cutting open cocoons to expose the pupae 
would probably have helped alleviate the problem. Peigler (1977) re- 
ported a similar problem with the emergence of C. promethea é x C. 
securifera (Maassen) 2 hybrids. 

One hybrid female was observed attempting to emit pheromone 


266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


around 2100 h. Her efforts were very brief and she deposited three or 
four infertile ova shortly afterward. Males confined in the same emer- 
gence cage did not respond to the female’s calling efforts. 

Several males were backcrossed to H. cecropia females. The males 
responded to the calling females during the latter’s normal mating time 
(0430 h to 0600 h), and the pairs remained coupled until early evening. 
Although females oviposited freely, no eggs hatched. Jim Tuttle (pers. 
comm.) observed a hybrid male respond to and mate with a calling C. 
angulifera female at 2200 h. The ova did not hatch. None of the hybrid 
males exhibited any difficulty clasping onto females. They apparently 
did not possess the shapeless valvae previously described, since such a 
developmental failure would probably be an obstacle to successful cop- 
ulation. 

It is of interest to compare the results of this study with those of 
Peigler’s (1978) C. angulifera 6 x S. cynthia 2 hybrid. Peigler experi- 
enced more difficulty throughout his study, as many of his larvae were 
lost to disease, several pupae died prior to emergence and no females 
were obtained. 

Collins and Weast (1961) obtained larvae from the cross H. cecropia 
6 x S. cynthia °. They stated, ““The larvae were raised to maturity on 
ailanthus and then lost.’’ The author has obtained several matings with 
S. cynthia as one parent and C. angulifera, C. promethea or H. ce- 
cropia as the other. Results have ranged from ova that did not hatch 
to larvae that failed to survive beyond the first instar. The less spec- 
tacular results of hybridization studies involving S. cynthia lend sup- 
port to the notion that Hyalophora and Callosamia are more closely 
related to one another than either genus is to the Asiatic Samia. Pop- 
ulations of S. cynthia in other faunal regions, including North America, 
are results of introductions by man. 

As noted in the introduction, no natural hybrids between Callosamia 
and Hyalophora are known. The natural mating times for Callosamia 
are mid-morning for C. securifera (Maassen), mid-afternoon for C. 
promethea and the hours preceding midnight for C. angulifera. Hya- 
lophora species mate in the hours immediately preceding dawn, thus, 
circadian mating behavior effectively eliminates intergeneric encoun- 
LErS. 

Examples of my hybrids are in my collection, in the collection of 
the Ohio Historical Society, Columbus, Ohio and in the United States 
National Museum of Natural History. 


ACKNOWLEDGMENTS 


I wish to thank Jim Tuttle, Troy, Michigan, for reviewing this paper, for his drawing 
and written description of the male genitalia, and for the photographs of the adult moths. 


VOLUME 38, NUMBER 4 267 


Glenn Firebaugh, Toledo, Ohio, photographed the mature larva. Thanks are also due 
Dr. Paul Brand, Toledo, and Dr. Paul M. Tuskes, Houston, Texas, for reviewing the text 
and offering many helpful suggestions. 


LITERATURE CITED 


CoLuins, M. M. & R. D. WEasT. 1961. Wild silk moths of the United States. Collins 
Radio Co., Cedar Rapids, Iowa. 138 pp. 

FERGUSON, D. C. 1972. The moths of America north of Mexico. Fasc. 20.2B, Bomby- 
coidea (in part). Classey, London. Pp. 155-269, pls. 12-22. 

PEIGLER, R. S. 1977. Hybridization of Callosamia (Saturniidae). J. Lepid. Soc. 31:23- 
34. 





1978. Hybrids between Callosamia and Samia (Saturniidae). J. Lepid. Soc. 32: 
191-197. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 268-280 


REVISION OF THE GENUS PARACHMA WALKER 
(PYRALIDAE: CHRYSAUGINAE) OF NORTH AMERICA 
NORTH OF MEXICO WITH DESCRIPTION OF A NEW GENUS 


EVERETT D. CASHATT 
Illinois State Museum, Springfield, Illinois 62706 


ABSTRACT. The genus Parachma Walker is redescribed. A lectotype is selected for 
P. borregalis Dyar, and P. borregalis is synonymized under P. ochracealis Walker. The 
color variation of P. ochracealis is discussed. A new genus, Basacallis, is described and 
P. tarachodes Dyar is designated the type species. Complete data and distribution maps 
are presented for all specimens examined. 


Twelve species have been placed in the genus Parachma Walker. 
Of these, two species, P. ochracealis Walker and P. borregalis Dyar 
were known from the United States and listed by Barnes and Mc- 
Dunnough (1917). A third species, P. tarachodes Dyar, listed in Hodges 
et al. (1983), was collected in the southeastern United States but re- 
mained unidentified in many collections. The present work, based on 
the accumulation of more specimens and intensive studies of genitalia, 
venation, and head characters, has resulted in a redescription of the 
genus Parachma. Parachma borregalis is treated as a synonym of P. 
ochracealis. Parachma tarachodes is removed from this genus and is 
designated as the type species for a new genus, Basacallis. 


Genus Parachma Walker, 1866 
(Figs. 1-5, 8-11) 


Parachma Walker, 1866:1263. Type species: Parachma ochracealis, by monotypy. 

Zazaca Walker, 1866:1269. Type species: Zazaca auratalis (=Parachma ochracealis 
Walker), by monotypy. 

Perseis Ragonot, 1891:538. Type species: Asopia culiculalis Hulst (=Parachma ochra- 
cealis Walker), by monotypy. 

Artopsis Dyar, 1908:95. Type species: Artopsis borregalis Dyar (=Parachma ochracealis 
Walker), by original description. 


Description 


Head. Labial palpus sharply upturned, about one-third longer than eye diameter, 
second segment about twice length of first and third segments; maxillary palpus vestigial, 
two segmented, hidden beneath scaling; proboscis well-developed, scaled at base; frons 
rounded, smoothly scaled; vertex smoothly scaled; antenna filiform, about seven-tenths 
forewing length, two rows of scales to each segment, uniformly pilose beneath; ocellus 
directly behind base of antenna; chaetosema formed by row of fine setae along ocular 
suture laterad and posterad to the ocellus. 

Thorax (Fig. 5). Forewing triangulate; Sc nearly straight, intercepting costa at about 
one-half length; R, short and arising just before anterior angle of discal cell; R,, Rs, Ry, 
and R, stalked together; R,, extremely short and weak, arising the same point as R, or 
short-stalked with R,; R, and R; stalked together, M, separate, arising from anterior angle 
of discal cell; M, and M, stalked together with Cu,; Cu, from below posterior of discal 
cell; 1A absent; 2A and 8A separate at base, anastomosed briefly a short distance from 


VOLUME 38, NUMBER 4 269 


the base then divergent; retinaculum normal. Hindwing with frenulum normal; Sc and 
Rs anastomosed beyond the discal cell; M, from anterior angle of discal cell; M, and M, 
coincident and stalked with Cu,, arising proximad to the posterior angle of the discal 
cell. Legs moderate in length; foreleg smoothly scaled; midtibia, midtarsi, hindtibia, 
hindtarsi with heavy scale tufts. 

Abdomen. Moderately long and slender. 

Male genitalia (Figs. 8, 9). Uncus moderately broad and hood-shaped, apex broadly 
rounded, lateral arms for articulation of gnathos relatively long; tegumen narrow dorsad; 
pedunculus strongly modified for articulation of gnathos; vinculum narrow ventrad, with- 
out a well-developed saccus; gnathos moderately long, not extending past apex of uncus; 
transtilla weak and incomplete; valva moderately developed with apex rounded, inner 
surface clothed with long setae directed dorsad; juxta relatively small and U-shaped; 
aedeagus moderately developed, cylindrical with base slightly broadened and without a 
caecum; vesica with microspines and two plate-like cornuti armed with a row of several 
spurs. 

Female genitalia (Fig. 10). Ovipositor moderately short; papillae anales moderately 
broad with apex unilobate; anterior apophysis broad at base, longer than posterior apoph- 
ysis; eighth segment relatively short; ostium bursae membranous and without a well- 
developed lamella postvaginalis; antrum relatively long and lightly sclerotized, constrict- 
ed at inception of ductus seminalis; ductus bursae broadened and sclerotized posteriad; 
anterior half narrowed and membranous; corpus bursae simple and without signa. 


Remarks 


Parachma is allied to Caphys Walker, 1863, Acallis Ragonot, 1891, 
and Zaboba Dyar, 1914c, as indicated by the venation and genitalia. 
All four genera have the forewing M, and M, long-stalked and Cu, 
separate, but in Parachma Cu, arises from the stem of M, and Ms. In 
all four genera, forewing 2A and 3A anastomose briefly as described 
above. The hindwing shows a similar relationship with M, and M; 
coincident and stalked with Cu, except in Acallis where Cu, is separate 
or arises from the same point on the discal cell. 

The male genitalia of Parachma, Acallis, Caphys, and Zaboba are 
similar in structure, having a simple well-developed valva, uncus, and 
gnathos. The uncus of Parachma is broad with a well-rounded apex, 
whereas in Caphys it is relatively narrow and tapered posteriad (Mun- 
roe, 1970, fig. 8). The uncus of Zaboba and Acallis is shorter than that 
of Parachma and more rounded than Caphys. 

The female genitalia are more diverse. Parachma and Acallis have 
an ovipositor that is moderate in length and without a lamella post- 
vaginalis. Caphys and Zaboba have an extended ovipositor with the 
lamella postvaginalis reduced to two small elongated plates. 


Parachma ochracealis Walker, 1866 


Parachma ochracealis Walker, 1866:12638. 

Zazaca auratalis Walker, 1866:1269. 

Asopia culiculalis Hulst, 1886:147. 

Artopsis borregalis Dyar, 1908:95. NEW SYNONYMY. 

Artopsis nua Dyar, 1914a:164. 

Parachma ochracealis a culiculalis, Barnes and McDunnough, 1917:138. 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-4. Wing maculation of Parachma ochracealis: 1, male, reddish brown form, 
Key Largo, FL; 2, female, reddish brown form, St. Petersburg, FL; 3, male, grayish 
brown form, Key Largo, FL; 4, female, pale ochreous form, Alpine, TX. 


Description 


Alar expanse. 13 to 23 mm. 

Head. Labial palpus ochreous with light to dark reddish brown laterad; frons and 
vertex ochreous, in darker specimens overscaled with dark reddish brown. 

Thorax. Dorsum light reddish brown to dark reddish brown or ochreous-gray, ventrum 
dark reddish brown to grayish brown. Forewing (Figs. 1-4) ochreous to reddish brown 
or ochreous-gray; antemedial line ochreous, extending from one-third costa excurved to 
about one-third hind margin; postmedial line ochreous, extending from about two-thirds 
costa to two-thirds hind margin, nearly straight; terminal line reddish brown; fringe 
ochreous; undersurface orange-brown to reddish brown, costa darker. Hindwing light to 
medium orange-brown; fringe ochreous; undersurface ochreous, overscaled with reddish 
brown to fuscous and with an ochreous median line. Legs dark reddish brown to fuscous; 
midtarsi ochreous, hind tibial spurs ochreous with band of dark reddish brown. 

Abdomen. Dorsum concolorous with wings, ventrum darker. 

Genitalia (Figs. 8-10). As described for the genus (Figs. 8-10). 

Type data. Parachma ochracealis Walker, holotype, male, no data, genitalia slide BM 
10729, in the collection of the British Museum (Natural History); Zazaca auratalis Walk- 
er, holotype, female, no data, genitalia slide BM 10730, in the collection of the British 
Museum (Natural History); Aspoia culiculalis Hulst, holotype, male, Florida (no other 
data), in the collection of the American Museum of Natural History; Artopsis borregalis 


VOLUME 38, NUMBER 4 Pri 





7 
? 


Fics. 5-7. Wing venation: 5, male, Parachma ochracealis: 6, male, Basacallis ta- 
rachodes; 7, female forewing. 


Dyar, four male syntypes [with identical labels], Los Borregos, Brownsville, Texas, June 
5, 1905, H. S. Barber (I hereby designate one of these syntypes as the lectotype of Artopsis 
borregalis and have so labeled it), U.S. National Museum Type No. 11921; Artopsis nua 
Dyar, holotype, male, Lakeland, Florida, March 1913, C. N. Ainslie, U.S. National Mu- 
seum Type No. 19081. 

Specimens examined (182 males (M), 109 females (F); Fig. 11). UNITED STATES, 
Arkansas: Hempstead Co., Hope, 12-VI-1926(1F), L. Knobal (CNC). Logan Co., Mt. 
Magazine, Cameron Bluff, 6-VI-1964(1F), 10-VI-1964(1M), gen. sls. EDC662, 765, J. F. 
G. & T. M. Clarke (USNM). Arizona: Cochise Co., Paradise, VI(1F), gen. sl. EDC759 
(USNM). Yavapai Co., Mayer, 18-VI-1959(1M), 25-VI-1959(2M), 11-VII-1959(1M), M. 
O. Glenn (INHS); 5 mi. N Prescott, 1-VII-1973(1F), 22-VI-1973(1F), elev. 5450 ft., L. 
M. Martin (LACM); 4 mi. N Prescott, 27-VI-1973(1F), 4 mi. N Prescott, Granite Dells, 
4-VII-1971(1M), L. M. Martin (LACM). Florida: Alachua Co., 3 mi. SW Gainesville, 
Archer Road Lab., 8-IV-1975(2M), J. B. Heppner (JBH), u.v. light; 9 mi. NW Gainesville, 
26/27-II-1975(1M), G. B. Fairchild (JBH), malaise trap; 9 mi. NW Gainesville, 5-IV- 
1975(1M), H. N. Greenbaum (JBH), malaise trap; Gainesville, 9-I[V-1963(1M), R. P. Esser 
(FSCA), 13-IV-1967(1M), R. P. Esser (FSCA); 18-IV-1968(1F), F. W. Mead (FSCA); 26- 
VII-1967(1M), J. W. Perry (FSCA), blacklight, 3-VI-1975(1M), D. H. Habeck (JBH), 
DHH rearing A1888, on media; Gainesville, Pine Hills Estates, 20-VIJ-1969(1M), H. V. 
Weems, Jr. (FSCA). Brevard Co., Cocoa Beach, Cape Kennedy, 9-VII-1967(1M), R. H. 
Leuschner (RHL); Cocoa Beach, Cape Kennedy, 17-VII-1967(1M), R. H. Leuschner 
(RHL). Charlotte Co., Punta Gorda, IIJ-1956(2M), 30-III-1941(1M), 14-IV-1956(1M,1F), 
25-IV-1941(1M), 15-80-IV-1941(1F), H. Ramstadt (INHS). Clay Co., Keystone Heights, 
6-ITI-1953(1M), H. E. Woodcock (FSCA), Lake Geneva, 16-III-1953(1M) H. E. Wood- 
cock (FSCA). Dade Co., Florida City, 25-III-1936(1M), gen. sl. EDC748, J. G. Francle- 
mont (CU); Homestead, 3-VII-1964(1M), 7-VII-1964(1M), 28-VII-1964(1M), 30-VII- 
1964(4M), 6-VIII-1964(1M), 13-VIII-1964(1M), 2-IX-1964(1M), 29-IX-1964(1F), D. O. 


272 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fics. 8-10. Genitalia of Parachma ochracealis: 8, male, ventral view, aedeagus re- 
moved, Benton Co., MO, slide EDC 1017; 9, aedeagus, lateral view of cornuti, Cochise 
Co., AZ, slide EDC 7583; 10, female, ventral view, spermatophore in corpus bursae, 
Benton Co., MO, slide EDC 1018. 


VOLUME 38, NUMBER 4 27 

















| 























Fic. 11. Documented locality records for Parachma ochracealis. One dot represents 
more than one locality where collection sites are adjacent. 


Wolfenbarger (MCZ). Hernando Co., Weeki Wachee Springs, 6-JJJ-1955(1F), 14-III- 
1955(38M,1F), 20-VII-1955(1M), 29-III-1955(1M,1F), J. F. May (MCZ), 14-ITI-1955(1F), 
J. F. May (CNC), 14-III-1955(1M), J. F. May (MCZ), 14-III-1955(1F), J. F. May (CNC), 
14-III-1955(1M), J. F. May (FSCA). Highlands Co., Archbold Biol. Stat., Lake Placid, 
15-31-VII-1948(1M), A. B. Klotts (AMNH), Lake Placid, VII-1945(1F), gen. sl. 749, J. 
G. Needham (CU). Liberty Co., Torreya State Park, 1-V-1952(1M), gen. sl. EDC666, J. 
R. McGillis (CNC); 1-V-1952(1M), O. Peck (CNC), 1-V-1952(1M), G. S. Walley (CNC). 
Manatee Co., Oneco, IV(2M), IV-1954(1M,1F), 5-V-1953(2M), 14-V-1953(1M), 6-VI- 
1953(1M,2F), 3-VII-1953(1F), 15-VII-1953(6M), 3-VIII-1953(1F), 27-VIII-1954(1F), 15- 
X-1953(1M), 30-X-1954(1F), P. Dillman (MCZ,CNC,FSCA). Marion Co., 12 mi. NW Salt 
Springs, Lake Delany Campground, u.v. light, 12-IV-1974(1M,2F), J. B. Heppner (JBH). 
Martin Co., Port Sewall, 7-11-II-1950(1M), 20-III-1938(1M), J. Sanford (AMNH). Monroe 
Co., Big Pine Key, 4-IX-1972(1M), at u.v. light, J. B. Heppner (JBH); Key Largo, 19-I- 
1967(1F), 26-I-1967(2M), 1-II-1965(1M,1F), 4-II-1967(1F), 10-IJ-1967(2M,1F), 11-II- 
1968(1F), Mrs. S. Kemp (LACM), 18-II-1967(1M,1F), 14-IJ-1967(1M), 21-II-1966(1F), 
27-II-1966(1F), 28-II-1967(1M), 11-III-1967(2M,1F), 14-IIJ-1967(1F), Mrs. S. Kemp (MCZ), 
31-III-1952(1M), G. S. Walley (CNC), 26-IV-1966(1F), 5-V-1966(1M), 12-V-1966(1F), 
22-VI-1966(1M), 17-VII-1965(1F), 25-VII-1966(1M), 22-VIII-1966(1M), Mrs. S. Kemp 
(MCZ); No Name Key, 20-IV-1974(1M,1F), at u.v. light, J. B. Heppner (JBH); Plantation 
Key, 4-IV-1966(3M), in blacklight trap, Zeigler & Weems (FSCA); Tavernier, 13-16- 
VIII-1955(2M,3F), J. N. Todd (CNC,FSCA). Okaloosa Co., Shalimar, 11-V-1964(3M), H. 
O. Hilton (FSCA). Orange Co., Orlando, 31-III-1942(1M), gen. sl. EDC767, D. F. Berry 
(CNC); Winter Park, VII-1946(1M), A. B. Klotts (AMNH). Pasco Co., 2 mi. E Land O 
Lakes, 4-VII-1973(7M,6F), E. D. Cashatt (ISM), live oak forest. Pinellas Co., St. Peters- 
burg, 12-IV-1915(1F), at light, gen. sl. EDC742, R. Ludwig (USNM); IV(1F), gen. sl. 
EDC110 (USNM). Polk Co., Lake Alfred, 5-VIJ-1928(1M,1F), gen. sl., EDC737,109, L. 
J. Bottimer (USNM). Putnam Co., 2 mi. S Welaka, 20-IV-1973(2F), at u.v. light, J. B. 
Heppner (JBH). Sarasota Co., no locality, 14-V-1946(1F), 25-V-1946(1M), C. P. Kimball 
(CNC,MCZ); Sarasota, 20-III-1946(1M), 26-III-1946(5M,2F), gen. sl. 744-8, 750-1, J. G. 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Needham (CU), 5-VII-1951(1M), at light, H. L. King (MCZ); Siesta Key, 30-II-1966(1M), 
30-III-1964(1M), 12-IV-1964(1M), 25-IV-1957(1M), C. P. Kimball (MCZ). Volusia Co., 
Cassadaga, 14-IV-1965(2M), 17-IV-1964(3M), 19-IV-1965(1M), 21-IV-1965(1M), 23-IV- 
1965(1F), 24-IV-1953(1F), 28-IV-1965(2F), 5-V-1961(1F), 6-V-1961(1F), 28-V-1962(1F), 
1-VI-1962(1F), 5-VII-1964(1F), 11-VII-1953(1M), 13-VII-1953(1M), 20-VII-1953(1M), 
2-VIII-1953(1M), 4-VIII-1955(1M), 7-VIII-1962(1F), 27-VII-1962(1F), 28-VIII-1962(1F), 
S. V. Fuller (FSCA). Walton Co., DeFuniak Springs, 13-IIJ-1958(1M), C. L. Dickenson 
(FSCA). No County Given, no date(1M,1F), P. Orange (INHS); V(1F), G. P. Hulst 
(AMNH); no date(3M), Acc. No. 26226, Mrs. A. J. Slosson (AMNH). Kentucky: Nelson 
Co., Leslie Farm, near Boston, 23-VI-1971(1M), G. Florence (UL). Missouri: Benton Co., 
4 mi. NW Warsaw, 6-VI-1970(4M,1F), 9-VI-1971(2F), gen. sl. EDC1018, u.v. light; near 
Warsaw, 3-VI-1967(1M), 3-VI-1971(1M), 9-VI-1971(1M), gen. sl. EDC1017, J. R. Heitz- 
man (JRH). Franklin Co., Meramac St. Park, 7-VI-1972(1F), J. R. Heitzman, old decid- 
uous forest (JRH). Mississippi: Forrest Co., Hattisburg, 29-V-1966(1F), Roshore (BM), 
VI-1944(1M), C. D. Michener (AMNH). Handcock Co., Big Biloxi, 12-V-1972(1M), 13- 
V-1973(1F), 15-V-1971(1M), 20-V-1972(1M), Kergosien (BM). Harrison Co., Bay St. Louis, 
1-VI-1978(2F), 2-VI-1971(1M), 3-VI-1971(1M), 4-VI-1979(1M,1F), 15-VI-1971(1M), 27- 
VI-1979(1M), 4-VII-1979(1F), 13-VII-1979(1M), 8-VIII-1971(1M), Kergosien (BM); 
Handsboro, 30-IV-1967(1M), Taylor (BM); Pass Christian, 1- VI-1979(2M), 25-VI-1979(1M), 
Kergosien (BM). Hinds Co., Clinton, 7-VI-1959(1M), 20-VI-1980(1F), B. Mather (BM), 
15-VI-1979(1F), Hartfield (BM). Jackson Co., Biloxi, 24-V-1964(1M), Taylor (BM), Ocean 
Springs, 27-V-1960(1M), 28-V-1960(1M), 30-VIII-1960(1F), K. Dawson (BM), 5-IX- 
1960(1M), gen. sl. EDC769, 9-IX-1960(1M), 17-IX-1960(1M), gen. sl. EDC768, K. Daw- 
son (CNC). North Carolina: Moore Co., Southern Pines, 8-VI-1915(1F), gen. sl. EDC760 
(USNM). New Mexico: Eddy Co., Whites City, 15-V-1950(1F), E. C. Johnson (CNC). 
Oklahoma: Cleveland Co., Norman, 22-V-1949(1F), W. J. Reinthal (CNC). Comanche 
Co., Wichita Mts., 6-V-1950(3M), 26-V-1950(1M), W. J. Reinthal (CNC). South Carolina: 
Charleston Co., 7 mi. NE McClellanville, Wedge Plantation, 6-VII-1973(2M,1F), E. D. 
Cashatt (ISM). Texas: Blanco Co., no locality, VIII(1F), G. D. Hulst (AMNH). Brewster 
Co., Alpine, 22-V-1950(2F), E. C. Johnston (CNC); Big Bend, 15-30-VII-1926(1M), O. 
C. Poling (CNC). Cameron Co., Brownsville, 20-IJJ-1908(1M), at light, no collector 
(USNM), 23-VIII-1931(1M), gen. sl. EDC734, T. N. Freeman (CNC), 7-XI(1M), gen. sl. 
EDC755 (USNM)); San Benito, 8-VIJ-1915(1M), gen. sl. EDC732 (USNM), 16-VII-1923(2F), 
gen. sl. EDC101 (USNM). Dallas Co., Irving, 2-V(1F), no collector (FSCA). Jeff Davis 
Co., Fort Davis, 20-V-1950(1M), E. C. Johnston (CNC); Limpia Canyon, 20-V-1950(6M), 
E. C. Johnston (CNC). Montague Co., no locality, 10-V-1941(1M), L. H. Bridwell (CNC). 
Nueces Co., Corpus Christi, 8-V-1943(1M), gen. sl. EDC735, W. M. Gordon (USNM), 
VII-1943(1M), at light, W. M. Gordon (CU). Walker Co., Stubblefield Lake, 12-V- 
1977(1M), u.v. light, Peigler & Brown (USNM). Zavalla Co., Nueces River, 28-IV-1910(1M), 
gen. sl. EDC102, F. C. Pratt (USNM). No County Given: X(1F), F. G. Schaup, gen. sl. 
EDC757 (USNM); IV(1M,1F), G. D. Hulst (AMNH); V(1M), G. D. Hulst (AMNH); 
VIK(1F), G. D. Hulst (AMNH); VIII(1M), G. D. Hulst (AMNH); no date(1M,6F), no 
collector (INHS). 

Life history. Dr. Dale H. Habeck, University of Florida, has reared larvae on a slightly 
modified Shorey and Hale (1965) pinto bean medium. The natural food habits and larval 
habits are unknown. 


Remarks 


The synonymy is a result of the wide range of color variation and 
size. The moths at hand indicate a clinal color relationship between 
the western and the southeastern specimens. Specimens from the 
warmer and more humid areas in Mississippi, Texas, and Florida are 
darker and more reddish. The lighter colored and larger moths were 
collected from Arizona, New Mexico, and western Texas (Fig. 4). Some 


VOLUME 388, NUMBER 4 275 


smaller specimens taken on the Gulf Coast are grayish brown (Fig. 3) 
and were described as Artopsis borregalis, a separate genus and species 
by Dyar (1908). The reddish form (Figs. 1, 2) was named culiculalis 
by Hulst (1886) and Artopsis nua by Dyar (1914a). One specimen from 
southern Florida has an ochreous forewing with a dark reddish brown 
median band. A few specimens exhibit variation in the arching and 
distance between the antemedial and postmedial lines. On examining 
a large series of specimens, I discovered a gradation of all of these 
characters. As Kimball (1965) so aptly stated, ““There is no question 
about the variation in color, as well as size, but it is difficult, if impos- 
sible, to fit specimens into the named forms.” I have examined all type 
specimens and have studied the genitalia. There are no apparent mor- 
phological differences between the ochreous, reddish brown, and gray- 
ish brown forms. 

This species comes in readily to a blacklight at night. The resting 
posture is distinctive, as for most of the Chrysauginae. The wings are 
positioned somewhat parallel to the substrate, not held roof-like over 
the abdomen. The legs are held at right angles from the body, dis- 
playing the heavy scale tufts on the mid and hindtibia. 


Genus Basacallis Cashatt, NEW GENUS 
(Figs. 6, 7, 10-17) 


Type species: Parachma tarachodes Dyar, 1914b. 


Description 


Head. Labial palpus curved upward, about one-fourth longer than eye diameter, first 
and third segments subequal in length, second about one-third longer; maxillary palpus 
vestigial, hidden beneath scaling; proboscis well-developed; frons rounded and smoothly 
scaled; vertex roughly scaled; antenna filiform, about seven-tenths length of forewing, 
each segment with two rows of scales, uniformly pilose beneath; ocellus posteriad to 
antenna base, chaetosema formed by a row of fine setae along ocular suture posteriad to 
ocellus. 

Thorax (Figs. 6, 7). Forewing long, relatively narrow and triangulate; Sc long; R, 
intercepts Sc in male, separate in female; R, and R, coincident and stalked with R,; male 
discal cell about one-third wing length, female discal cell about one-half wing length; 
M, separate, arising from anterior angle of discal cell; M,, Ms, Cu,, and Cu, all separate 
and arising from posterior angle of discal cell in male, Cu, arising proximad to posterior 
angle in female; 2A and 8A separate at base, briefly anastomosed a short distance from 
base; retinaculum normal. Hind wing with frenulum normal; Sc and Rs anastomosed 
beyond discal cell; discal cell extremely short with posterior angle long; M, coincident 
with M, and arising from the posterior angle of the discal cell; Cu, and Cu, separate. 
Legs moderately long; midtibia with two scale tufts. 

Abdomen. Moderately long with no lateral scale tufts. 

Male genitalia (Figs. 14, 15). Uncus relatively short, tapered posteriad, lateral arms 
strongly modified for articulation with gnathos; tegumen narrow dorsad; pedunculus 
strongly modified for articulation with gnathos; vinculum moderately broad; gnathos 
long, slender and aculeate with apex hooked dorsad; valva moderately long and wide 
with tip curved mediad, sacculus slightly expanded, transtilla weak; juxta narrower at 
base; aedeagus long and slender, distal one-third bent ventrad. 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 12, 18. Wing maculation of Basacallis tarachodes: 12, male; 13, female; both 
from Pensacola, FL. 


Female genitalia (Fig. 16). Ovipositor relatively short; apex of papillae anales unilo- 
bate; anterior and posterior apophysis slender and moderately long, approximately the 
same length; lamella postvaginalis triangulate with anterior margin cleft; ostium bursae 
relatively small and membranous; ductus bursae extremely slender. 


Remarks 


The type species of Basacallis, tarachodes Dyar, is not congeneric 
with P. ochracealis, but belongs in a separate genus allied to Humiphila 
Becker (1974), a genus described for a saprophagus species (H. paleo- 
livacea Becker) in Costa Rica. The male and female genitalia show 
relatively close relationships between H. paleolivacea and B. tara- 
chodes in the gross morphology. In particular, the aedeagus (Fig. 15) 
is acutely bent in these two species. 

Of the North American fauna Basacallis shows the closest natural 
relationships with Caphys and Acallis. Hindwing M, and M, are coin- 
cident and the discal cell is reduced. Vein M, + M;, and Cu, are short- 
stalked or arise separately from the posterior angle of the discal cell as 
in Caphys and Acallis. Basacallis is distinguished from the related 
genera by forewing M, and M, arising separately and not stalked as in 
Caphys and Acallis. The male genitalia more closely resemble Acallis 
except for the acutely bent aedeagus. The bursa copulatrix is relatively 
slender and delicate in Acallis and Basacallis except for the scleroti- 
zation of the ductus bursae just below the inception of the ductus 
seminalis. I have found the structure of the bursa copulatrix to be 
extremely delicate and difficult to dissect and interpret. 

The name Basacallis is feminine in gender and formed by combin- 
ing the Greek word for foundation and the genus name Acallis (Bas 
+ acallis). 


VOLUME 388, NUMBER 4 aT 





lmm 





Fics. 14-16. Genitalia of Basacallis tarachodes: 14, male, ventral view, aedeagus 
removed; 15, aedeagus, slide EDC 872; 16, female, ventral view, slide EDC 873; all 
from Pensacola, FL. 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 














@ Basacallis tarachodes (Dyar) 























Fic. 17. Documented locality records for Basacallis tarachodes. One dot represents 
more than one locality where collection sites are adjacent. 


Basacallis tarachodes (Dyar), NEW COMBINATION 


Parachma tarachodes Dyar, 1914b:306. 


Description 


Alar expanse. 16 to 23 mm. 

Head. Labial palpus gray, brownish laterad; frons, vertex, and antenna gray. 

Thorax. Dorsum gray, ventrum reddish-brown. Forewing (Figs. 12, 18) light gray, 
with darker gray median band across middle one-third, antemedial and postmedial near- 
ly straight and distinct in male, more diffuse and slightly excurved in female; terminal 
line fuscous; fringe reddish to purplish brown, sometimes invading distal portions and 
fading proximad; underside purplish red with short fuscous antemedial and postmedial 
lines from the costa. Hindwing grayish white with outer margin purplish, fading proxi- 
mad, terminal line fuscous; fringe ochreous with base reddish brown; females darker, 
underside reddish to purplish brown with an ochreous median band. Legs reddish brown 
to fuscous, scale tufts fuscous, midtarsi ochreous; hindtibia and tarsi ochreous overscaled 
with fuscous. 

Abdomen. Dorsum ochreous-gray, ventrum light reddish brown. 

Genitalia (Figs. 14-16). As described for the genus. 

Type data. Holotype, female, Portobelo, Panama, April 1912, genitalia slide EDC878, 
Type No. 16297, in the U.S. National Museum. 

Material examined (8 males (M), 6 females (F); Fig. 17). UNITED STATES, Florida: 
Escambia Co., Pensacola, III-1961(1M), 12-IV-1962(1F), 19-IV-1961(1M), 20-IV-1961(1M), 
gen. & wing sl. EDC872, 20-VIII-1961(1F), gen. & wing sl. EDC873, S. Hills (MCZ). 
Manatee Co., Oneco, IV-1954(1M), 14-V-1953(1F), V-1954(1F), gen. sl. EDC878, P. 
Dillman (CNC). Mississippi: Harrison Co., Handsboro, 16-IV-1966(1M), Taylor (BM). 
Warren Co., Bovina, 14-IV-1972(1M), 26-VII-1976(1M), 21-IX-1972(1M); Vicksburg, 3-IX- 


VOLUME 38, NUMBER 4 279 


1981(1F), 20-IX-1978(1F), B. Mather (BM). South Carolina: Charleston Co., Mc- 
Clellanville, Wedge Plantation, 3-V-1973(1M), at light, R. B. Dominick (RBD). 
Life history. Unknown. 


Remarks 


Unrecognized in many collections, this species was referred to in 
Kimball (1965) as, “5801,3 [X.] SP. — either Xantippe, Parachma, or 
a closely related genus.”’ Of the few specimens examined, the size and 
maculation varies considerably between males and females. The alar 
expanse ranges from 16 to 18 mm in males and 19 to 23 mm in females. 
The forewing median band is sharply defined by distinct antemedial 
and postmedial lines in the male. In females the antemedial and post- 
medial lines are less distinct and the median band is darker than that 
of the male. 


ACKNOWLEDGMENTS 


I am grateful to the following persons and/or institutions for their patience, assistance 
and the loan of their specimens, without which this study would not have been possible: 
American Museum of Natural History (AMNH); British Museum (Natural History), Bryant 
Mather (BM); Canadian National Collection (CNC); Cornell University (CU); Charles V. 
Covell, Jr., Dale H. Habeck, Florida State Collection of Arthropods (FSCA)); Illinois 
Natural History Survey (INHS); Illinois State Museum (ISM); John B. Heppner (JBH); J. 
R. Heitzman (JRH); Museum of Comparative Zoology (MCZ); Richard B. Dominick 
(RBD); Ronald H. Leushner (RHL); University of Louisville (UL); United States National 
Museum (USNM). Genitalia photographs were taken by Scott Kilborne, Southern Illinois 
University Medical School. Thanks go to James R. Purdue, my associate, for his assistance 
with the word processor and to George L. Godfrey, Illinois Natural History Survey, for 
his review of the manuscript. 


LITERATURE CITED 


BARNES, W., & J. MCDUNNOUGH. 1917. Check list of the Lepidoptera of boreal Amer- 
ica. Herald Press, Decatur, Ill. IX + 392 pp. 

BECKER, V. O. 1974. Studies on the shootborer Hypsipyla grandella (Zeller) (Lepidop- 
tera, Pyralidae). XXVI. A new genus and three new species of Microlepidoptera 
(Pyralidae and Grazillariidae) associated with Carapa, Cedrela, and Swietenia in 
Costa Rica. Turrialba 24(3):332-335. 

Dyark, H.G. 1908. A review of the North American Chrysauginae. Proc. Entomol. Soc. 
Washington 10:92-96. 

1914a. New American Lepidoptera. Insecutor Inscitiae Menstruus 2:161-164. 

1914b. Report on the Lepidoptera of the Smithsonian Biological Survey of the 

Panama Canal Zone. Proc. U.S. Natl. Mus. 47:139-350. 

1914c. Descriptions of new species and genera of Lepidoptera from Mexico. 
Proc. U.S. Natl. Mus. 47:365-409. 

HOpGESs, R. W. (Editor). 1983. Checklist of the Lepidoptera of North America North 
of Mexico. E. W. Classey Limited and The Wedge Entomological Research Foun- 
dation, London, England. 284 pp. 

HULST, G. D. 1886. Descriptions of new Pyralidae. Trans. American Entomol. Soc. 13: 
145-168. 

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Arthropods of Florida and neigh- 
boring land areas 1:v + 363 pp., 26 pls. 











280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


MUNROE, E. 1970. A new genus and three new species of Chrysauginae (Lepidoptera: 
Pyralidae). Can. Entomol. 102:414—420. 

RAGONOT, E. L. 1890 [1891]. Essai sur la classification des Pyralites. Ann. Soc. Ent. 
France (6)10:436-546. 

SHOREY, H. H. & R. L. HALE. 1965. Mass-rearing of the larvae of nine noctuid species 
on a simple artificial medium. J. Econ. Entomol. 58(3):522-524. 

WALKER, F. 1863. List of the specimens of lepidopterous insects in the collection of 
the British Museum. London. Part 27:1-286. 

1865 [1866]. List of the specimens of lepidopterous insects in the collection of 

the British Museum. London. Part 34:1121-1534. 





Journal of the Lepidopterists’ Society 
38(4), 1984, 281-309 


THE BIOLOGY AND DISTRIBUTION OF CALIFORNIA 
HEMILEUCINAE (SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge 141G, Houston, Texas 77054 


ABSTRACT. The distribution, biology, and larval host plants for the 14 species and 
subspecies of California Hemileucinae are discussed in detail. In addition, the immature 
stages of Hemileuca neumogeni and Coloradia velda are described for the first time. 
The relationships among the Hemileuca are examined with respect to six species groups, 
based on adult and larval characters, host plant relationships and pheromone interactions. 
The tricolor, eglanterina, and nevadensis groups are more distinctive than the electra, 
burnsi, or diana groups, but all are closely related. Species groups are used to exemplify 
evolutionary trends within this large but cohesive genus. 


The saturniid fauna of the western United States is dominated by 
moths of the tribe Hemileucinae. Three genera in this tribe commonly 
occur north of Mexico: Hemileuca, Coloradia, and Automeris. Al- 
though no Automeris are native to California about 50% of the Hemi- 
leuca and Coloradia species in the United States occur in the state. 
The absence of Automeris and other species from California is due to 
the state’s effective isolation from southern Arizona and mainland Mex- 
ico by harsh mountains, deserts, the Gulf of California, and climatic 
differences. The Hemileuca of northern Arizona, Nevada, and Utah 
are very similar to that of California, while those of Oregon, Washing- 
ton, and Idaho represent subsets of the northern California fauna. 

The majority of the saturniid species in the United States have had 
little or no impact on man, but some Hemileucinae have been of eco- 
nomic importance. In California, the larvae and pupae of Coloradia 
pandora have been utilized by indians as a food source (Aldrich, 1911, 
1921). Other species, especially in the genera Automeris, Hemileuca 
and Saturnia have urticating spines which can inflict a painful sting; 
the resulting welt can persist for days. Larvae of the range caterpillar, 
Hemileuca oliviae, have often reached pest status by damaging range 
grasses utilized by cattle in Oklahoma and Texas (Ainslie, 1910; Watts 
& Everett, 1976; Huddleston et al., 1976; Coleman, 1982). The larvae 
of Coloradia pandora undergo periodic outbreaks at which time the 
larvae have damaged or defoliated large stands of pines (Chamberlin, 
1922; Patterson, 1929; Wygant, 1941). In California, pandora reaches 
the status of a pest about once every 30 years. Other species have been 
nuisances locally on crops or ornamentals but have never consistently 
been abundant or caused economic losses. 

In the last 30 years there have been two major publications dealing 
with the Saturniidae of the United States. In his work on the Saturni- 
idae of the Western Hemisphere Michener (1952) dealt with the mor- 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


phology, phylogeny, and classification of the group. Ferguson (1971, 
1972) authored a two part series on the Saturniidae of the United States 
and Canada and illustrated in color most of the species, presented line 
drawings of the male genitalia, and summarized their biology. Fer- 
guson’s work condensed most of the available information on the sa- 
turniid fauna north of Mexico. It became clear that relatively little was 
known about the distribution, biology, and immature stages of many 
western species, particularly in the tribe Hemileucinae. The purpose 
of this paper is to discuss the Hemileucinae of California and to present 
new information regarding their biology, distribution, and immature 
stages. 

Unless specifically indicated, diagnostic information on adults or im- 
mature stages is not intended as redescriptions but merely to help the 
reader recognize the uniqueness of each taxon. Distribution records are 
based on a review of specimens in the collections of: The University 
of California at Berkeley, Davis, and Riverside; California State Uni- 
versities at San Diego, San Jose, Fresno, and Humboldt; Natural His- 
tory Museums of San Diego, Los Angeles, and Santa Barbara as well 
as the collection at the California Academy of Science, San Francisco; 
the private collections of Steve McElfresh, Ken Hansen, John Johnson, 
Mike Collins, Sterling Mattoon and the author. In addition the author 
has traveled and collected extensively in this area and has reared all 
of the species occurring in the western states. Information on flight 
period is based on observation and capture records for wild specimens 
in collections; emergence dates from specimens which were obviously 
reared were excluded, as was distribution data on mislabeled speci- 
mens. Distribution maps are provided to show trends, and in most cases 
the maps should be considered as a conservative estimate (Fig. 5). 


Genus Hemileuca Walker 


Hemileuca are medium to large moths with wingspans ranging from 
3 to 9 cm. Moths in this genus include nocturnal and diurnal species 
and occur from desert to alpine habitat. Although some species are 
widely distributed with ranges from Mexico to Canada, the majority 
are restricted to Arizona, California, Nevada, Texas, and Mexico. Eight 
species of Hemileuca occur in California representing four of the six 
species groups. Michener (1952) recognized four subgenera and treated 
Pseudohazis as a junior synonym of Hemileuca based on adult mor- 
phology. Ferguson (1971) and Tuskes (1978) presented additional in- 
formation which supports the merger of Pseudohazis and Hemileuca. 
Hemileuca chinatiensis (Tinkham) and H. griffini Tuskes (Fig. 2m) 
have genitalia, wing shapes, wingspans, sexual dimorphism and larval 
characters which place them at a transitional point between Hemileuca 


VOLUME 38, NUMBER 4 283 





Fic. 1. Hemileuca of California. a & b. H. eglanterina eglanterina, $ & 2; ¢ & d, 
H. eglanterina shastaensis; e, H. eglanterina eglanterina form denudata; f & g, H. 
nuttalli uniformis; h, H. hera hera; i, H. eglanterina annulata; j, H. eglanterina an- 
nulata, Elko Co., Nevada; k, H. hera marcata, Klam. Co., Oregon; 1 & o, H. hera hera, 
Modoc Co., Calif. 


and Pseudohazis (Table 1). Within the genus there are six distinct 
species groups which are based on male genitalia, wing pattern, exter- 
nal adult morphology, behavior, larval morphology, and larval host 
plant preferences at the family level (Table 1). 


284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Hemileuca and Coloradia of California. a, H. electra electra 2; b, H. electra 
electra 6, e, H. juno 2; d, H. juno 6; e, H. neumoegeni 2; f, H. burnsi 2; g, H. electra 
clio 9°; h, H. electra clio 6; i, H. nevadensis 2; j, H. nevadensis 6, k, H. neumoegeni 4; 1, 
H. burnsi 6; m, H. griffini 6, Az.; n, H. diana 4, Az.; o, H. tricolor 6, Az.; p, C. velda 4; 
q, C. velda 9; r, C. pandora lindseyi 6. Hemileuca griffini, H. diana, and H. tricolor do 
not occur in California but were included as representatives of species groups or transi- 
tional taxa discussed in the text; the latter three species are from Arizona. 


Pheromone attraction tests show positive interaction between many 
species groups (Table 2). For example, female electra will attract and 
mate with males of burnsi, diana, and eglanterina as well as all eglan- 
terina subspecies. Females of nevadensis, which are superficially sim- 


VOLUME 38, NUMBER 4 285 


TABLE 1. Relationships among the Hemileuca of the United States. 
2? much Larval Pupation 
Male genitalia Time of larger paniculum _in soil/ 
grouping* Primary hostplant oviposition than 6 concolor leaf litter 
electra group 
electra Polygonaceae day/night yes no yes 
juno Leguminosae day/night yes no yes 
burnsi group 
burnsi Compositae/ Rosaceae night yes no yes 
neumoegeni Rosaceae /Anarcadi- night yes no yes 
aceae 
diana group 
diana Fagaceae day yes no yes 
grotei Fagaceae day yes no yes 
maia group 
nevadensis Salicaceae day yes no yes 
maia Fagaceae/Salicaceae day yes no yes 
lucina Rosaceae day yes no yes 
eglanterina group 
eglanterina Rosaceae/Salicaceae/ day no yes yes 
Celastraceae 
nuttalli Caprifoliaceae/Rosa- day no yes yes 
ceae 
hera Compositae day no yes yes 
griffini Rosaceae day some some yes 
chinatiensis Leguminosae day yes no yes 
tricolor group 
tricolor Leguminosae night yes no no 
oliviae Gramineae night yes no no 
hualapai Gramineae night yes no no 


* Wing pattern grouping is the same. 


ilar to those of the diana group, will not attract either diana or grotei 
males. Based on Tables 1 and 2, the tricolor, eglanterina and neva- 
densis groups are the most distinctive, and any further revision which 
might reestablish Pseudohazis to generic status would have to recog- 
nize the tricolor and perhaps the nevadensis groups as unique. Such 
splitting would accomplish little, and hide the morphological, biolog- 
ical, and behavioral relationships which they share. Much needs to be 
learned about the Mexican species before the status of the entire group 
can be dealt with properly. Presently the Hemileuca appear to be a 
rather cohesive group containing distinctive but closely related assem- 
blages of species. 


Biology of Hemileuca 


Adult moths emerge from their pupation site in the soil or leaf litter 
during the morning and climb shrubs or grasses prior to wing expan- 


286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Intrageneric attraction and mating between 11 western species of Hemi- 
leuca. Unless otherwise indicated tests were made with caged females and wild males in 


the field. 


Calling Results Calling Results 


female Male tested At./Ma. female Male tested At./Ma. 
burnsi electra no* electra eglanterina yes 
chinatiensis diana no* annulata 
chinatiensis eglanterina no electra eglanterina yes 
chinatiensis nevadensis no* shastaensis 
chinatiensis eglanterina no electra hera no 
shastaensis electra nuttalli no/yes* 
diana nevadensis no* electra burnsi yes 
diana grotei yes electra diana yes 
diana juno no? electra electra clio yes 
eglanterina nuttalli yes hera eglanterina no 
eglanterina hera no hera nuttalli no 
eglanterina eglanterina yes juno diana yes 
shastaensis nevadensis grotei no 
eglanterina eglanterina yes nevadensis lucina yes* 
annulata nuttalli hera no 
eglanterina nevadensis no* nuttalli eglanterina no/yes* 
electra eglanterina yes 


At./Ma. = attraction and mating. 
* — male reared and caged with female. 


sion. Newly emerged females release pheromones that may attract 
dozens of males to their location within a few minutes. Usually females 
begin to emit pheromones after their wings have expanded. Mating 
requires 20 minutes to an hour, during which time the pair remain 
almost motionless. In captivity, male eglanterina have mated consec- 
utively with three females, each of which produced fertile ova. 

Shortly after mating the female deposits ova in a ring (Fig. 3k) 
around the branch she is perched on (if it is the host plant) or may fly 
for a short period of time prior to oviposition. After the first egg ring 
is completed she frequently flies for five to 15 minutes and then de- 
posits a second egg ring which usually contains fewer eggs than the 
first. Depending on the species, eight to over 300 ova may be deposited 
in a ring. Oviposition generally occurs on stems 2 to 8 mm in diameter. 
Although females of our western species usually mate only once, in 
certain populations of H. maia the female may mate again after de- 
positing the first egg ring. 

The majority of eggs in any given ring usually hatch within two 
days of each other. Early instar larvae are black, but as they mature, 
species specific color patterns develop. Since many of the western species 
hatch between December and April, the dark coloration and their 
gregarious nature may play an important role in their thermoregula- 
tion (Fig. 3l). Larval phenotypes of eglanterina, nuttalli, griffini, and 


VOLUME 38, NUMBER 4 287 


bie ape 





Fic. 3. Last instar Hemileuca larvae of California. a, H. eglanterina shastaensis; b, 
H. eglanterina annulata; ec. H. nuttalli uniformis; d, H. nuttalli nuttalli; e, H. hera 
hera; f, H. neumoegeni; g, H. burnsi; h, H. electra electra; i, H. juno; j, H. nevadensis; 
k, H. nevadensis egg ring; 1, First instar larvae of H. nevadensis feeding on willow. 


nevadensis are generally consistent within a population, but striking 
differences are often found between different populations (Figs. 3a, b, 
c, d). The larvae of electra, burnsi, juno, neumoegeni, and hera usually 
exhibit little variability. A key to the last instar saturniid larvae of the 
West Coast has been published, and includes some host plant and 
habitat data (Tuskes, 1976). All Hemileuca larvae are covered with 


288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Cc} a 
ik PA. \ / ‘a 


Fic. 4. Coloradia larvae of California. a, C. velda, mature; b, C. velda, mature; e, 
C. velda (4th instar); d, C. pandora lindseyi, mature blackish phenotype (Mono Co., 
Ca.); e, C. pandora lindseyi, mature brown phenotype (San Diego Co., Ca.); f, C. pan- 
dora lindseyi egg cluster on pine needle. 


short secondary setae and various types of urticating scoli on the dorsal 
and lateral surfaces. When disturbed gregarious larvae may respond 
by moving the body segments anterior to the prolegs back and forth 
at a frequency of about 1 cycle per second. At times almost every larva 
in the cluster may oscillate in synchrony. The urticating scoli of Hemi- 
leuca, Automeris, and Saturnia larvae are capable of inflicting a severe 
sting. The irritation may last for half an hour, and the affected area 
may discolor; welts lasting from one to 14 days may develop. The 
intensity of the pain inflicted by the larval scoli seems to vary from 
one species to the next. 

In the 4th or 5th instar the larvae lose their gregarious tendencies 
and feed individually. At this time they disperse over a wider area of 
the host plant or to surrounding plants. When a larva is mature it leaves 
the host plant and wanders on the ground in search of a suitable lo- 
cation to pupate. All of the California species pupate either in the leaf 
litter, under objects such as rocks, burrow into the ground, or utilize 
deep cracks in the soil. The pupal cell consists of debris or soil held 
together with a loose matrix of silk which forms a cup over the pupa 
with little or no silk below. The cell is fragile and merely picking it 
up often results in its destruction. An exception are the members of 


VOLUME 38, NUMBER 4 289 


the tricolor group, many of which spin a loose but complete silken 
cocoon that is attached to vegetation. 

The length of time spent in the pupal stage for any given species 
varies from population to population. In areas with a long growing 
season, overwintering eggs hatch in the spring, develop, and emerge 
as adults later that same summer. Where the growing season is short 
(e.g. coastal fog belt, or subalpine) adults and larvae from different 
generations may overlap. Pupae from spring larvae may emerge, mate, 
and deposit eggs, or overwinter and hatch the following summer. Thus, 
in some populations, eggs and pupae rather than just eggs function as 
the overwintering stage. I refer to this as an asynchronous two year 
life cycle with an adult flight each year. All of the California species 
have the ability to spend two or more years in the pupal stage. 


Hemileuca eglanterina eglanterina (Boisduval) 


Hemileuca eglanterina (Figs. la, b, e) is the only western saturniid 
with a dorsal forewing that is yellow, black, pink, and occasionally 
beige. The antemedial and postmedial lines and margin are black. In 
some populations the females have more pink on the forewing than 
males. Dorsally, the hindwings are black and yellow. The ventral wing 
surface is always black, yellow, beige, and frequently pink. The ab- 
domen of both sexes is alternately banded black and yellow, but in 
some individuals one color may dominate the pattern. There is a great 
deal of geographic variation within this species. Specimens from south- 
ern California are often smaller and have less pink on the forewings 
than northern populations. The form “‘denudata”’ is widespread in pop- 
ulations from central California to Washington, although the frequen- 
cy of its occurrence varies greatly from one population to the next. All 
specimens expressing the “denudata”’ trait have been males in which 
the black markings are either greatly reduced or absent, therefore the 
wings are predominantly pink or yellow. Another distinctive pheno- 
type is one in which the yellow has been replaced by brown. 

Typical eglanterina blends into the Great Basin subspecies, annu- 
lata, along the passes on the east slopes of the Sierra Nevada. A second 
subspecies, shastaensis is characterized by melanic adults and occurs 
in extreme northern California and southern Oregon. Male and female 
eglanterina are of similar size. California specimens of the nominate 
form have wingspans that range from 63 to 72 mm. 

The flight season extends from late June to early October, but the 
majority of the records are from mid-July through the last week of 
August. Both males and females are active day flyers. 

Hemileuca eglanterina occurs in a wide variety of habitats. In dry 
areas such as the Central Valley, eglanterina occurs in the riparian 


290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


habitat where the larval host plants are various species of Salix. In the 
chaparral community, as well as in pine, oak, or redwood forests, the 
larvae feed extensively on Ceanothus and Cercocarpus. Host plants 
which are used less frequently or by specific populations include: Pru- 
nus, Pyrocantha, Purshia, Rubus, Quercus, and Symphoricarpos. On 
occasion this species has been common in residential areas and or- 
chards. 

Although the phenotype of the last instar larva is fairly uniform 
within a given population, there is a great deal of variability between 
populations. Last instar larvae have 1 to 3 complete cream colored 
lateral lines, but in some populations they are absent. The ventral 
surface, intersegmental area, and prolegs vary in coloration from black 
to red. The ground color is black, dark gray, or dark brown; the dorsal 
scoli are yellow and black. 

Egg rings on Salix are easily located during the winter. Each ring 
may consist of 75 to over 250 ova. In the Central Valley the eggs begin 
to hatch in mid to late March, and last instar larvae can be collected 
from mid-May to mid-June. At the lower elevations eglanterina has a 
one year life cycle, but at higher elevations or along the coast it has 
an asynchronous two year life cycle with an adult flight each year. 


Hemileuca eglanterina shastaensis (Grote) 


Hemileuca eglanterina shastaensis (Figs. 1c, d) is distinguished from 
typical eglanterina by its melanic tendencies and distribution. Al- 
though some males are totally black on the dorsal surface, the majority 
have a moderate amount of pink on the forewing and yellow on the 
hindwing. Even the darkest males express 3 of the 4 basic eglanterina 
colors on the ventral wing surface: black, yellow, and pink. Males 
exhibit a full range of phenotypes from black to only slightly melanic. 
Females have a definite tendency to be lighter in coloration, and some 
appear similar to the nominate form except for having a more exten- 
sive rose-pink coloration on the dorsal forewing. Specimens in Califor- 
nia collections have wingspans which range from 65 to 78 mm. 

The flight season extends from mid-June to mid-August, but the 
majority of the records are from mid-July. Both males and females are 
active day flyers and are usually closely associated with the chaparral 
plant community. 

Larval masses have been collected by the author numerous times on 
Purshia tridentata (Pursh) in California and Oregon and reared to 
maturity in captivity on Cercocarpus. Wild cherry is another possible 
host according to Ferguson (1971). All of these plants are in the family 
Rosaceae, a group commonly used by typical eglanterina. Early instar 
larvae of shastaensis may be collected in May or June, and mature 


VOLUME 38, NUMBER 4 291 


larvae looking for pupation sites are frequently observed crossing roads 
in mid-July to August. Mature larvae measure 50 to 70 mm in length 
and are reddish brown with bold cream colored lateral lines; the in- 
tersegmental area is deep red. This subspecies has a two year asyn- 
chronous life cycle on Mt. Shasta as adults and larvae can be collected 
at the same time. Larvae reared in the Central Valley exhibited a one 
year life cycle (Fig. 3a). 

A cross between a female shastaensis and a typical male eglanterina 
produced offspring that appeared almost identical to the male parent, 
except that they had a deep rose color on the forewing which is char- 
acteristic of many shastaensis. Because of staggered emergence the F, 
could not be backcrossed or selfed. Additional matings between shas- 
taensis and nominate eglanterina are needed to determine their re- 
lationship to each other. 


Hemileuca eglanterina annulata Ferguson 


Hemileuca eglanterina annulata (Figs. 1h, i) has the typical eglan- 
terina pattern but may be distinguished from the nominate form by 
the reduction or absence of pink on the forewing and a tendency for 
the black markings to be more extensive, often with a smudged ap- 
pearance. In California, specimens of annulata may be confused with 
H. nuttalli uniformis, but the two species may be separated by ex- 
amining the hindwing postmedial black line between vein M, and the 
inner margin of the wing; this line is straight or convex in annulata 
and concave in nuttalli. In addition, the ventral surface of California 
annulata is usually black, yellow, and beige, while California nuttalli 
are black and yellow. Some annulata females are only black and yel- 
low; if so, the forewing postmedial coloration will separate them from 
female nuttalli. In Nevada and Arizona, annulata is larger than nut- 
talli. California specimens have wingspans that range from 60 to 78 
mm. Specimens from Nevada and Arizona reach 85 mm and are fre- 
quently darker. 

The flight season of annulata extends from late June to early Sep- 
tember, with the majority of the records from mid-July to mid-August. 
Both males and females are active day flyers from 1000 to 1630 h, 
with a peak near 1200 h. 

In California, annulata is restricted to the Great Basin habitat on 
the dry eastern slopes of the Sierra Nevada, where it integrates with 
the nominate form. Specimens from this area are the same size as the 
nominate form but usually have smudged black maculation and lack 
bold pink markings on the forewings. Some individuals have a slightly 
diffuse pinkish yellow cast over the forewing, while other populations 
have little or no pink. Although California populations are variable and 


292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


are both smaller and lighter in coloration than Nevada and Arizona 
populations, they are best described as annulata with some genetic 
input from typical eglanterina. The adults are usually seen flying near 
the larval host plants which include: Prunus emarginata (Dougl.), Ce- 
anothus velutinus (Doug].), Purshia tridentata (Pursh), and Sympho- 
ricarpos vaccinoides (Rydb.). Mature larvae from California are sim- 
ilar to those from the Central Valley except the cream colored lateral 
stripes are more prominent, and the ventral surface is usually light red 


(Fig. 3b). 


Hemileuca nuttalli uniformis (Cockerell) 


The forewing of male nuttalli uniformis (Figs. lf, g) is cream to 
beige dorsally, with a black basal patch, antemedial, and postmedial 
line, discal spot, and margin. In addition (as in eglanterina and hera), 
black wedge-shaped marks extend from the margin towards the post- 
medial line. The forewing of the female is very similar to the male, 
but the area between the margin and the postmedial line is yellow in 
the female rather than beige as in the male. The ventral surface of the 
fore- and hindwings are yellow and black. The thoracic tufts are yel- 
low, and the abdomen has alternate bands of black and yellow. This 
species is frequently sympatric with annulata, and the two are often 
confused. For diagnostic differences see H. eglanterina annulata. Male 
and female nuttalli are of similar size; wingspans range from 60 to 68 
mm. Specimens from Arizona and Nevada are frequently larger, 66 to 
78 mm, and their pattern is more diffuse. 

In California, nuttalli flies from mid-July to perhaps mid-Septem- 
ber, with the peak flight period in August. Both males and females are 
active day flyers. 

This subspecies inhabits the eastern slopes of the Sierra Nevada from 
Inyo Co. north to Alpine Co. at elevations generally ranging from 2150 
to 3400 m. Although probably widespread in this area, limited access 
to the proper habitat has resulted in relatively few collecting records. 
Specimens from Nevada and northern Arizona are slightly larger, and 
the black markings are frequently more diffused than Sierran material. 
Ferguson (1971) suggested that the uniformis phenotype may be the 
result of its cool environment. On two occasions the author reared 
Sierran uniformis in the hot Central Valley, and the resulting adults 
expressed the uwniformis phenotype. 

Although frequently sympatric with H. eglanterina annulata and 
H. hera in the mountains, uniformis infrequently occurs on the high 
plateau east of the Sierra, an area dominated by sagebrush and saline 
soils. Collins and Tuskes (1979) found that male wniformis are attracted 
to and will mate with female eglanterina. Such interspecific matings 


VOLUME 38, NUMBER 4 293 


are minimized in the wild by a partially allochronic flight period. 
Female uniformis have a tendency to mate later in the day than fe- 
male eglanterina. Male uniformis fly from 1030 to 1800 h, with 75% 
of their flight activity after 1330 h. Male eglanterina annulata at the 
same location flew from 1030 to 1630 h, with about 75% of their flight 
activity prior to 1830 h. Female uniformis did not attract male an- 
nulata. 

Overwintering egg rings hatch in late April or May and pupation 
occurs in July. The larval host plants in California are Purshia triden- 
tata (Pursh) and Symphoricarpos vaccinoides (Rydb.). In the Sierra 
Nevada uniformis has an asynchronous two year life cycle with an 
adult flight each year. When reared at lower elevations uniformis has 
a one year cycle. The larvae of uniformis from Monitor Pass, Mono 
Co. have 1 prominent and 2 poorly developed cream colored lateral 
lines. The ground color is black and there are numerous grayish short 
secondary setae; the dorsal rosette setae are yellow and black. The head 
is black and mature larvae measure 53 to 63 mm in length (Fig. 3c). 


Hemileuca nuttalli nuttalli (Stecker) 


Adult nuttalli nuttalli difter from nuttalli uniformis in subtle ways. 
The nominate form is said to be larger and the black markings well 
defined rather than diffused as in uniformis (Ferguson, 1971). In ad- 
dition, the forewing of female nuttalli has a uniform yellow cast with 
black maculation, while that of uniformis is beige with a yellow cast 
only between the postmedial line and wing margin; the maculation is 
also black but slightly more diffused. 

Flight data were not available since all of the California records 
have been from reared specimens. In Oregon this form flies during 
July and August. Both males and females are active day flyers. 

Typical nuttalli is common in southern Oregon and has been col- 
lected in extreme northern California. In Siskiyou Co. the larvae have 
been found feeding on Purshia tridentata and Symphoricarpos. Pu- 
pation occurs during July and early August; adults emerged the follow- 
ing August. The larvae of nuttalli are extemely variable. MacFarland 
(1974) described mature nominate nuttalli from eastern Oregon as 
black, with no maculation, and with black scoli. Larvae from Siskiyou 
Co., CA have prominent yellow dorsal scoli, and 3 distinct cream col- 
ored lateral lines; the ground color is black, and there are numerous 
short secondary setae on the segmental areas (Fig. 3d). 


Hemileuca hera hera (Harris) 


The fore- and hindwing of H. hera (Fig. 1h) are white with a black 
basal patch, antemedial line, postmedial line, discal spot, and marginal 


294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


areas. The costal area is black, and the veins between the marginal and 
postmedial line are covered with triangular-shaped patches of black 
scales. The head and anterior portion of the thorax are yellow, dorsal 
thoracic area black, thoracic tufts white. The abdomen is alternately 
banded with yellow, black, and occasionally white. Specimens from 
the Sierra Nevada are frequently darker than those from the adjoining 
Great Basin. Both male and female hera are of similar size; the wing- 
span of California specimens ranges from 59 to 73 mm. 

The flight season extends from late June to mid-September with the 
majority of the records from July to early August. Both males and 
females are active day flyers. 

The larvae of hera feed almost exclusively on one species of sage, 
Artemisia tridentata Nutt. (Basin sagebrush), but when larval popu- 
lations are dense they have been found occasionally feeding on various 
Lupinus and Eriogonum. Although A. tridentata is widespread in 
California, hera occurs only in the Great Basin habitat on the east side 
of the Sierra-Cascade ranges; from Inyo Co. north to Modoc Co., and 
is common in adjoining areas of Nevada and eastern Oregon. 

Some populations of hera in northern California exhibit a great deal 
of variation. A series from Davis Creek, Modoc Co. contained typical 
specimens, melanic individuals, and the form marcata which has been 
treated as a subspecies (Figs. 1l-o). The form “marcata”’ (Fig. 1k) is 
one in which the black markings on the wings are greatly reduced; 
this form may be similar to “denudata” in H. eglanterina populations. 
The extent of variation observed in the Davis Creek area is almost 
identical to that found in nearby populations of H. eglanterina shas- 
taensis. Further study is needed to determine the status of “marcata”’ 
and the extent of variation in the southern Oregon populations. Hemi- 
leuca hera magnifica (Rotger) is a subspecies from Colorado and New 
Mexico for which little biological data are available. Larvae from both 
states were collected and reared on A. tridentata. The larvae of mag- 
nifica were virtually identical in color and pattern to nominate hera 
larvae from Monitor Pass, Mono Co., California (Fig. 3e). 

In California, overwintering eggs hatch in late April or May. Early 
instar larvae are black and gregarious; pupation occurs during July and 
August. Populations express a two year asynchronous life cycle with 
adults flying each year. When reared at lower elevations the moth 
expresses a one year life cycle, although some may hold over in the 
pupal stage for up to two years. McFarland (1974) published a partial 
description of a mature larva. A complete description of a mature 5th 
instar larva from Mono Pass, Mono Co., California is presented (Fig. 
3e). 


VOLUME 38, NUMBER 4 295 


Description of Last Instar Larva 


Head: Shiny black with numerous white secondary setae; diameter, 4.5-5.0 mm. Clyp- 
eus black with 4 long setae. Body: Length 55 to 62 mm; width, 9 mm. Ground color 
black. Ventral surface, light gray intersegmentally and gray-brown to black elsewhere. 
Sublateral scoli black. Lateral scoli with yellow rosette and branching black stalk. Dorsal 
scoli from thoracic segment 3 to abdominal segment 8 yellow rosette type. Body covered 
with numerous white secondary setae. Dorsal and lateral surfaces black. Body with 3 
distinct lateral and 1 dorsal cream-white colored lines. Line I most conspicuous, passing 
through the sublateral scoli and extending length of larva. Line II, slightly ventral to 
lateral scoli and best developed in intersegmental area. Line III, midway between lateral 
and dorsal scoli, appearing as series of dashes. Line IV consists of series of dots or lines 
in intersegmental area and anterior portion of each segment paralleling black middorsal 
line. Thoracic shield, anal shield, and lateral shields of prolegs black. True legs black. 
Spiracles orange. 


Hemileuca electra electra Wright 


The dorsal forewing surface of electra (Figs. 2a, b) has a white 
medial area; the discal spot, costa, postmedial line, and margins are 
black. Although the forewing postmedial line is present in the females, 
its occurrence is variable in the males. The ventral surface of the fore- 
wing and both surfaces of the hindwings are red and black. The thorax 
is gray to black with white thoracic tufts and collar. The abdomen is 
red dorsally and black and white ventrally. Hemileuca electra can be 
distinguished from the Mojave Desert subspecies clio, by the well de- 
veloped black postmedial line on the forewing of nominate females 
and the white area or series of white patches that occur between the 
postmedial line and the black wing margin. The latter characteristic is 
usually but not always diagnostic. In male electra the dark postmedial 
line or smudged area on the forewing touches the discal spot, whereas, 
in California clio populations it does not; male and female clio lack 
the dark postmedial line. The wingspan of male electra ranges from 
45 to 54 mm; the females range from 53 to 62 mm. 

Hemileuca electra flies from late August to early December with 
the majority of the records from mid-September to early October. Both 
sexes are active day flyers. 

Nominate electra is restricted to the coastal chaparral plant com- 
munity of five southern California counties and portions of northern 
Baja California, Mexico. Males have a fast erratic flight except when 
approaching a female. Adults fly from about 1000 to 1500 h and may 
be collected as they perch on various chaparral plants in the evening. 
Although wild females are infrequently captured, larvae and egg rings 
are frequently locally abundant. 

The egg rings are deposited in the fall on the floral stems of Eriog- 
onum fasciculatum Benth. Each egg ring contains 30 to 60 greenish 
ova which hatch in December or January. The host plant grows com- 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


monly in disturbed habitats such as road cuts and hill slides. In non- 
disturbed areas it is frequently associated with Rhus laurina and coastal 
sages. Mature larvae are found in late March or April. Comstock and 
Damme rs (1939) described the mature larva and reported its length to 
be 45 mm, but most mature field collected larvae are larger. The larva 
is black with many secondary white setae extending from white pi- 
nacula giving it a gray-brown appearance. There are three continuous 
lateral cream-yellow colored lines extending the length of the body. 
The intersegmental area is red, and the setae of the dorsal rosette are 
yellow and black (Fig. 3h). 


Hemileuca electra clio Barnes & McDunnough 


Hemileuca electra clio (Figs. 2g—h) is similar in coloration to typical 
electra, but there are major differences in the pattern of the forewing. 
In clio from California the white medial patch of the forewing is well- 
defined and the postmedial line is undefined. Male and female clio are 
very similar in appearance. Specimens of clio from Arizona are usually 
larger, darker, and have a slightly different flight period than Califor- 
nia populations. Some of the Arizona populations are almost completely 
black dorsally except for the basal area of the hindwing which is red. 
The wingspan of male clio from California ranges from 51 to 59 mm; 
females 51 to 69 mm. 

California records indicate the flight period of clio is shorter than 
typical electra. Collecting records range from the first week of Septem- 
ber to the first week of November with the peak flight period occurring 
in mid-September. Male clio are active day flyers, while females are 
usually captured in the afternoon and have been collected numerous 
times during the early evening at blacklights. 

Hemileuca electra clio is found in the high desert regions of southern 
California and is frequently sympatric with H. burnsi and H. neu- 
moegeni. The area occupied by clio has less precipitation, hotter sum- 
mers and colder winters than the coastal chaparral community occu- 
pied by typical electra. Some populations of clio (in Riverside, San 
Bernardino, Imperial, and San Diego counties) on the eastern passes of 
the mountains which separate the chaparral from the desert plant com- 
munity have hybrid like phenotypes suggesting that gene exchange has 
or does occur between the two taxa. 

The author has crossed nominate electra from San Diego Co. with 
a wild male electra clio from Phelan, San Bernardino Co. The F, 
females were extremely variable, but definitely expressed the clio phe- 
notype. The smallest female had a wingspan of 53 mm, was infertile, 
and had melanic forewings. The other females had melanic tendencies 
and wingspans that ranged from 63 to 68 mm. The males from this 


VOLUME 38, NUMBER 4 297 


cross were similar in appearance to, but on the average (54-58 mm) 
slightly larger than, normal clio. 

The egg rings of clio contain 7 to 26 large gray eggs. The ova of 
clio are a different color, larger, and the rings contain far fewer eggs 
that those of typical electra. The larvae of clio feed on Eriogonum 
fasciculatum var. polifolium and are distinguished from typical elec- 
tra by their larger size and coloration. Last instar larvae of clio from 
California and Arizona are similar in appearance; they have fewer 
white pinacula than electra electra and are darker in appearance. In 
addition the dorsal rosettes are solid black in clio but black and yellow 
in typical electra. Smith (1974) also noted these and other differences 
between the larvae. 


Hemileuca juno Packard 


The forewing of juno (Figs. 2c, d) is black with a white medial area. 
The hindwing of the male is black except for the center of the discal 
spot which is white. Females usually have lightly marked white medial 
area on the hindwings. Some specimens, especially females, have a 
gray-brown rather than black wing coloration. The thorax of the male 
is black while that of the female is frequently gray. The abdomen of 
the male is black anteriorly and red posteriorly; females are black with 
a red tip. The pattern of juno is similar to that of electra clio from 
Arizona. The wingspan of male juno from Arizona ranges from 52 to 
59 mm; females 58 to 64 mm. 

Although juno is widespread from Idaho to northern Mexico (Com- 
stock and Dammers, 1939), it is infrequently collected except in Ari- 
zona and southern New Mexico. Hemileuca juno is assumed to occur 
in southern California, but the only record is from San Diego (Co.) and 
was collected in 1908. The larval host plant, Prosopis juliflora DC. 
(mesquite), grows commonly in the area of Anza-Borrego and many 
other locations in southern California. Despite a great deal of collecting 
activity in this area, there are no recent reports of larvae or adults from 
the California desert. The moth has been collected on the California-— 
Arizona border at Yuma (XI-10-64). 

In Arizona, first instar larvae are found in late April or early May. 
Although larvae will feed on the leaves of the host plant, they prefer 
the buds and flowers which develop in clusters. In captivity some larvae 
refuse to-feed on anything but the flowers or buds. Pupation occurs in 
leaf litter near the surface of the ground during late June. 

The mature larva measures about 50 mm in length and is black with 
numerous secondary setae extending from white pinacula which gives 
the larva an overall gray appearance. The head and true legs are black 
and the prolegs and ventral surface are gray-brown. The dorsal rosette 


298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


setae are uniquely colored with red at the base and black at the tips 
of the spines. There are 3 white and 1 black lines, all of which appear 
as a series of dashes on the lateral surface and extend the length of the 
larva. Comstock and Dammers (1989) described and illustrated the 
mature larva of juno, but their description was not complete (Fig. 3i). 

Adults have been captured from late September to early December, 
with the majority of the records from later October to early November. 
Emergence occurs in the morning, and males begin to fly between 
1000 and 11380 h. During a peak flight in 1982 females were observed 
ovipositing from late afternoon to at least sunset. The male flight 
dropped off considerably by 1530 h, and shortly thereafter, males were 
found perched on the host plant. By 1700 h most moths that were 
observed flying were females; although uncommon, females have been 
collected at blacklights. 


Hemileuca burnsi Watson 


The wings of H. burnsi (Figs. 2f, |) are chalk white and black. The 
antemedial black line is solid and is never intersected by an elongated 
spot as in H. neumoegeni. The black discal spot appears as a ring on 
the forewing but on the hindwing is often reduced to a simple black 
dot. The postmedial band is well developed on the forewing but fre- 
quently reduced or absent from the hindwing of males. Although the 
abdomen of the female is usually white, it may have semicircular bands 
of red and/or black; the abdomen of the male is white. The wingspan 
of male burnsi ranges from 48 to 52 mm; females 58 to 62 mm. 

Collection dates indicate that the flight season extends from the last 
week of August to the first week of November, with the majority of 
the records from the second and third weeks of September. Records 
from Arizona, Nevada, and Utah also indicate September to be the 
month of the peak flight period in those states. 

Male burnsi are active day flyers, generally flying between 0900 and 
1500 h. They seek virgin females that remain perched on desert shrubs 
and transmit pheromone during the day. Females oviposit at night and 
are strongly attracted to light for the first few hours after it becomes 
dark and less frequently through the rest of the night. Upon emergence 
males are chalk white, but with age some become cream colored. Older 
specimens in collections are frequently light yellow. 

The eggs overwinter and usually hatch in late February or early 
March. Early instar larvae are black and gregarious. Late instars are 
black with white pinacula, giving them a gray appearance. In addition 
the mature larva has at least two cream colored lateral lines extending 
the length of the body. Pupation occurs in April or early May. Com- 


VOLUME 38, NUMBER 4 299 


stock and Dammers (1937) described the first and last instar larva as 
well as the pupa of burnsi (Fig. 8g). 

In the Reno area of Nevada the larval host plants are Tetradymia 
glabrata Gray and Dalea fremontii Torr. In southern California they 
utilize Tetradymia axillaris Nels. = T. spinosa H. & A., Prunus fas- 
ciculata (Gray), and various species of Dalea. Other plants associated 
with the burnsi habitat on the higher desert slopes near Little Rock 
and Phelan include: Artemisia, Chrysothamnus, Ephedra, Eriogonum 
fasciculatum var. polifolium (Benth.), Larrea, Lycium, Purshia, and 
Yucca. Although late instar larvae have been collected on Eriogonum 
fasciculatum var. polifolium, I have found that early instar larvae 
cannot survive on that host. After the 4th instar, successful develop- 
ment on Eriogonum did occur, and the resulting adults were of normal 
size. 

Hemileuca burnsi is sympatric with H. electra clio in many areas 
of Kern, Los Angeles, Riverside and San Bernardino counties. I have 
seen one male specimen which may represent a hybrid between these 
two species. On occasion male burnsi have been attracted to a calling 
female of H. electra. 


Hemileuca neumoegeni (Henry Edwards) 


The wings of H. neumoegeni (Figs. 2e, k) are lustrous white. The 
distal spot of the fore- and hindwings is black and sickle-shaped with 
white centers; the antemedial and postmedial lines are also black. An 
elliptical black spot with a white center may occur in the antemedial 
forewing line. The thorax and head are red and the thoracic tufts are 
white. The abdomen of both sexes are similarly marked; red dorsally 
and laterally with a red and white ventral surface. Some females have 
white scaling on the dorsal posterior portion of the abdomen. Hemi- 
leuca neumoegeni is occasionally confused with H. burnsi, but the two 
are easily separated. The wings of burnsi are chalk-white and the 
antemedial line on the forewing is not interrupted by an elongated 
spot. The abdomen of burnsi adults is primarily or completely white 
rather than red as in neumoegeni. Male neumoegeni have a wingspan 
of 48 to 55 mm; the females from 58 to 64 mm. 

The flight season extends from the second week of August to early 
October, with the majority of the records from the second and third 
week of September. Arizona, Nevada, and Utah records also indicate 
September as the month when most specimens have been captured. In 
California it appears that the occurrence of warm thunder storms early 
in the flight season enhances the September flight, while cold fronts 
have a retarding effect. Both males and females are nocturnal flyers 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and are attracted to lights; most specimens have been collected prior 
to 2230 h. 

The distribution of this species is more extensive than previously 
indicated (Ferguson, 1971); for in addition to Arizona, it has been 
collected in Utah, Nevada, and California. California records date back 
to the early 1930’s. Adults and larvae have been collected in the Clark, 
Granite, Ivanpah, Mescal, Mid Hills, New York and Providence Moun- 
tain ranges of San Bernardino Co. 

Adults emerge during the late morning but remain inactive during 
the day; mating occurs after twilight. There is some variation among 
the males with regard to the size and presence or absence of an orbic- 
ular-like spot in the antemedial line. Perhaps 20% of the California 
specimens lack or have a very reduced orbicular-like spot. 

Females deposit 15 to 30 grayish ova in a ring near the terminal 
ends of the host plant and then move to another plant before depositing 
the next egg ring. In the Providence and New York Mts., egg rings 
and larvae have been collected numerous times on Prunus fasciculata 
(Torr.) and occasionally on Rhus trilobata anisophylla (Greene). Lar- 
vae in the Spring Mts. of Nevada were found on P. fasciculata by Eric 
Walter, while Mike Van Buskirk collected larvae south of Flagstaff, 
Arizona on R. trilobata. Both host plants are associated with dry slopes 
and washes in the Mojave and Colorado deserts above 1000 m. Bauer 
(1948) reported Eriogonum fasciculatum Benth. as the host plant and 
was cited by Ferguson (1971), but this record appears to be incorrect 
and may have resulted from misidentified larvae. 

In California the ova hatch between early April and May. During 
the early instars the larvae are black and feed gregariously. In captivity 
young larvae on potted plants fed randomly throughout the day, but 
Ath and 5th instar larvae descended the secondary stems and remained 
on the main stems, frequently near the base of the plant during the 
day, and fed on the leaves of the terminal branches at night. Because 
mature neumoegeni larvae have a reddish brown intersegmental area 
and may be gray rather than black, they resemble H. electra larvae 
rather than those of H. burnsi, to which it is most closely related. 
Pupation occurs during June. 

Hemileuca neumoegeni is sympatric with Hemileuca electra clio B. 
& McD. in the Providence and New York Mts., and populations of H. 
burnsi Watson occur in these same mountains. The larvae of electra 
clio feed on Eriogonum, primarily fasciculatum, while burnsi larvae 
feed on Dalea, Tetradymia, and Prunus fasciculata. Because of dif- 
ferential mating times, morning vs evening, there is little probability 
of interaction between neumoegeni and burnsi or electra. The larvae 
of neumoegeni are described here for the first time (Fig. 3f). 


VOLUME 38, NUMBER 4 301 


Description of Larval Instars 


FIRST INSTAR. Head: Black, diameter 0.9 mm. Body: Length 6 mm, width 1.2 mm. 
Ground color, solid black. All scoli except dorsal thoracic and dorsal lateral simple with 
one long seta extending from each. Dorsal thoracic and dorsal lateral scoli forked at tip, 
with one slightly elongated seta extending from each branch. All scoli black. 

SECOND INSTAR. Head: Black, diameter 1.4-1.5 mm. Body: Length 10-11 mm, width 
2.4 mm. Ground color solid black. All scoli branched and black with black spines. True 
legs, prolegs, and spiracles black. 

THIRD INSTAR. Head: Black, diameter 2.2-2.4 mm. Body: Length 17 mm, width 4.1 
mm. Ground color solid black. Larva similar to second instar only larger. 

FOURTH INSTAR. Head: Black, diameter 2.8-3.1 mm. Body: Length 24-86 mm. Ground 
color black. Ventral surface black with some white setae. Lateral surface with single 
undulating continuous white line extending length of larva and passing through base of 
lateral scoli. Faint traces of second white lateral line appears in segmental area only, in 
line with dorsal lateral scoli. Segmental area with numerous short secondary setae, some 
of which extend from cream to light pink pinacula. Intersegmental areas black. Mid 
dorsal area black. All scoli black and branched type. Prolegs black and covered with 
short white secondary setae. Planta orange. Spiracles black. 

FIFTH INSTAR. Head: Black with numerous short white secondary setae, diameter 4— 
4.6 mm. Clypeus black. Body: Length 52-57 mm, width 10-11 mm. Ground color black 
to gray. Ventral surface gray with short gray secondary setae extending from white 
pinacula; intersegmental area red to flesh colored. Ventral and sublateral scoli with black 
shafts and white or black spines. Lateral scoli with black shafts with black and white 
spines and numerous white and black spines at base of shaft. Dorsal scoli on thoracic (T) 
segment 8 to abdominal (A) segment 8 of rosette type with gray centers and black tips. 
Three white or cream colored lateral lines divide lateral and dorsal surfaces into distinct 
color regions. Line I continuous and well developed undulating white-gray line extending 
length of larva, passing through base of sublateral scoli. Line II appears as series of gray 
dashes in intersegmental area only, below line of lateral scoli. Lateral surface between 
line I and II black with secondary setae extending from gray pinacula, giving area gray 
appearance. Intersegmental area with fewer pinacula and appearing black. Line III 
poorly developed and interrupted by reddish brown intersegmental area. Line III just 
ventral of dorsal scoli and extending length of larva. Segmental area between line II and 
III black; but secondary setae extending from gray pinacula giving area gray appearance; 
intersegmental area brick-red. Gray-black segmental and red intersegmental pattern 
continuing to black mid dorsal line. True legs black. Prolegs gray with light brown to 
red planta. Dorsal thoracic shield black. Proleg shield light brown to red-brown. Spiracles 
orange. 


Hemileuca nevadensis Stretch 


The wings of H. nevadensis (Figs. 2i, j) are black to dark gray with 
a wide cream colored medial band; both fore- and hindwings are sim- 
ilarly marked. The black discal spot on the forewing is lunate and 
larger than that of the hindwing. The abdomen is black; males have a 
distinctive red tuft at the tip, while females have a black or whitish 
tip. Some minor geographical variation is observed, as specimens from 
Nevada and northeastern California have slightly narrower black mar- 
ginal borders than specimens from more southerly areas of California. 
The wingspan of California specimens ranges from 53 to 66 mm for 
the males; 58 to 67 mm for females. 

The flight season extends from the third week of September to early 
December, with the majority of the records from the second week of 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


October through the first week of November. Both males and females 
are active day flyers. 

Hemileuca nevadensis is widespread and locally abundant from ex- 
treme northern Baja California to the Los Angeles-Riverside basin and 
through the Central Valley to Turlock. In California this species is 
closely associated with the riparian habitat, where it feeds on Salix. 
The same host plant and habitat is occupied by H. eglanterina in the 
northern part of the Central Valley. This may have some influence on 
the distribution of nevadensis. There are scattered records for neva- 
densis along the Colorado River and the east side of the Sierras and 
Cascade ranges. Some populations on the east side of the Sierras may 
feed on Populus in addition to Salix. 

The adults emerge in the morning and usually mate prior to noon. 
On cool cloudy days, moths have been seen in large numbers perched 
on plants along creek and river banks. Females may deposit over 200 
ova in a ring on the woody thin stems of willow (Fig. 3k). The ova 
overwinter and hatch in late March or April. The immature stages 
have been described by Wright (1888), Dyar (1895), Packard (1914), 
and Comstock and Dammers (1989). Early instar larvae are black (Fig. 
31), but as the larva matures it develops yellow colored pinacula and 
assumes a black and yellow ground color. The prolegs have reddish 
brown patches near their base, and the prothoracic and anal shields 
are red. Larvae from northeastern California and Nevada have more 
yellow markings than those from southern or central California (Figs. 


3j, 1). 
Genus Coloradia Blake 


Adult Coloradia are characterized by their nocturnal habits, mod- 
erate to large size and their gray to black cryptic forewings. The 
hindwings are gray and either pink, cream, or light brown. Male an- 
tennae are quadripectinate, while those of the female are bipectinate 
or biserrate. Unlike Hemileuca the labial palps are large, separate, and 
not fused. Four species of Coloradia occur north of Mexico, two of 
which occur in California. 

Members of the genus are widespread in pine forests of the western 
United States and portions of Mexico. The larvae feed on various species 
of pine. Coloradia pandora and its subspecies have caused economic 
damage to ponderosa, Jeffrey, and lodgepole pine forests in the western 
United States (Patterson, 1929; Wygant, 1941). Some species have a 
one year life cycle, while others usually require two years to develop 
from the egg to the adult. The larvae of Coloradia are similar to those 
of Hemileuca but have a dark gray to brown ground color and gen- 
erally lack well developed cream or white lateral lines. In addition 


VOLUME 38, NUMBER 4 303 


some Coloradia lack the finely branched or rosette type scoli, and 
generally the spines are not urticating. 


Coloradia pandora lindseyi Barnes & Benjamin 


Male and female C. pandora (Fig. 2r) are similar in appearance but 
differ in size. Dorsally, the forewings vary from gray-brown to dark 
gray. The antemedial and postmedial lines are black and trimmed with 
white; the submarginal band is light gray. The hindwing marginal area 
is brown to dark gray, while the submarginal portion is white to pink. 
There is a brown or dark gray postmedial band and a distinct black 
discal spot. The remainder of the hindwing is cream, gray, or pink. 
The head, abdomen, and thorax are dark gray to brown with tufts of 
white scales. In California, pandora could be confused with C. velda, 
but the two are easily separated by size, flight season and appearance. 
The hindwing postmedial line of velda is diffused and frequently 
touches the discal spot, and the majority of the wing is gray; only 20 
to 35% of the basal area is pink. In pandora the hindwing postmedial 
line is well defined and does not touch the discal spot; the majority of 
the hindwing is light brown, cream or pink. In California the wingspan 
of male pandora varies from 65 to 76 mm; females from 74 to 95 mm. 

The flight season extends from mid-July to early October, with the 
majority of the records from early August to mid-September. Both 
sexes are nocturnal flyers and are attracted to lights. During massive 
population buildups, moths may fly during the day. 

Coloradia pandora lindseyi inhabits Jeffrey pine forests and at times 
may be extremely abundant. Females deposit clusters of large white 
ova on the branches or needles of the pines (Fig. 4f). The eggs hatch 
in September or October, and the larvae begin to feed on the old 
needles. They pass the winter in the early instars high in the trees. 
Mature larvae descend the trees in June and July and pupate in the 
soil. The pupae may hatch that year or overwinter and hatch the 
following summer. Mature larvae measure over 58 mm in length, and 
have a brown to brown-black ground color (Figs. 4d, e). The posterior 
portion of each segment is dark brown, while the anterior and inter- 
segmental area is light brown. There is a thin black mid dorsal line 
surrounded on each side by a thick dorsal white line. Two white lateral 
lines extend the length of the larva. The body is covered with light 
brown to cream colored pinacula from which extend short brownish 
secondary setae. The head is light brown. All scoli are short and the 
branches are much less developed than those of Hemileuca larvae (Fig. 
4d). Although not described in detail, Patterson (1929) illustrated the 
immature stages of lindseyi. 

Aldrich (1911, 1921) found that the Piute Indians near Mono Lake, 


304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Mono Co. utilized the mature larvae as a food source, which they called 
pe-aggie. Once collected, the larvae were killed by heating, then dried 
and stored for future use. Engelhardt (1924) reported that indians in 
southern Oregon also utilized lindseyi larvae as a food source. 

Mature larvae collected in the Laguna Mts. of San Diego Co. during 
early June pupated in loose soil. The pupation chamber was cup-shaped 
and consisted of soil and debris held together with a small amount of 
silk. Some adults emerged two months later in August, while others 
emerged in August of the following year. Similar results were observed 
by Aldrich (1921) with larvae collected at Mono Lake. Carolin (1971) 
found that lindseyi may remain in the pupal stage at least six years 
prior to emergence, but the majority emerged as adults during the first 
two years. 

Populations of pandora undergo periodic outbreaks, during which 
time larval densities are sufficiently high to cause serious damage to 
commercial stands of pine. Coloradia pandora lindseyi damaged Pinus 
ponderosa Laws. in Oregon (Chamberlin, 1922; Patterson, 1929), but 
seems to prefer Pinus jeffreyi Grev. & Balf. in California. During the 
1976 to 1982 outbreak near Mammoth Lakes, Mono Co., the larvae 
fed primarily on Jeffery pine but on occasion were found feeding on 
ponderosa and lodgepole pines (P. contorta Dougl.). Various papers 
have dealt with the biology and economic impact of lindseyi (Cham- 
berlin, 1922; Engelhardt, 1924; Aldrich, 1921) but only Patterson (1929) 
provides much detailed information. Patterson did state that lindseyi 
is strictly a diurnal species, an observation which would distinguish this 
subspecies from other Coloradia (Ferguson, 1971). Contrary to Patter- 
son’s observations, I have collected lindseyi commonly at lights, and 
generally after 2100 h. Chamberlin (1922) apparently worked with 
Patterson during the 1919 Klamath Falls, Oregon outbreak and implies 
that the moths were attracted to lights. During the Mono Co. outbreak, 
the U.S. Forest Service estimates that 5000 acres of Jeffrey pine were 
defoliated in 1979 and about 16,000 acres in 1981 (Schaaf, 1980, 1981). 
Although the U.S. Forest Service only mentioned the nocturnal flights 
of the moth in their environmental assessment report, John Johnson 
informed me that some adults were observed flying during the day. It 
appears that the behavior of this species may change when populations 
are at high densities. During population explosions there is a potential 
for day flights, although most will probably still be active at night. At 
normal population levels all flight activity (mate seeking and oviposi- 
tion) is at night. 


Coloradia velda Johnson and Walter 


The color and pattern of male and female Coloradia velda (Figs. 
2p, q) are similar in appearance. The forewings are gray to gray- 


VOLUME 38, NUMBER 4 305 


brown; the antemedial and postmedial lines are dark gray to black and 
accented with white scales. The forewing discal spot is darker gray, 
smaller, and better defined than that of the hindwing. The hindwings 
are primarily gray to gray-brown with a slightly darker postmedial 
line that is adjoined to a distal pink line. The basal % to % of the wing 
is pink, as is the inner margin of the wing. In some specimens the 
hindwings appear partially transparent. The thorax and abdomen are 
dark gray with white scales mixed throughout. The only species similar 
to velda in California is C. pandora lindseyi, which is a larger species 
that flies later in the year. The hindwing of pandora is predominantly 
white or white and pink, and the discal spot is free of the gray post- 
medial line. In velda the gray from the margin extends all the way to 
the discal spot, and there are no extensive white areas. For additional 
comments see C. pandora lindseyi. Male velda have wingspans ranging 
from 51 to 61 mm; females from 58 to 73 mm. 

Specimens have been captured from May to the end of July, but the 
majority of the records are for June. The adults are nocturnal and 
attracted to lights. 

Presently velda is known only from the mountains of San Bernardino 
County. This species was recently described as a distinct species by 
Johnson and Walter (1980). Prior to its description it had been assumed 
to be a uniquely disjunct population of C. doris Barns. 

Only the life history of C. pandora and its various subspecies have 
been described in the literature. Based on this information, the life 
cycle of velda is notably different from that of pandora. Larvae of 
velda emerge from the egg in early July and feed exclusively on pinyon 
pine, Pinus monophylla Torr. & Frem. Attempts to rear them from 
the Ist instar on P. jeffreyi were unsuccessful, but mature larvae may 
accept alternate hosts. The larvae feed gregariously in the early instars 
and singly in the 4th and 5th. They develop rapidly and after five 
instars pupate in mid-September. Larvae enter loose soil and burrow 
to a depth of 10 to 15 cm, then construct a structure around them by 
attaching silk to large soil particles and plant debris. The resulting 
pupae overwinter and adults emerge the following summer. By con- 
trast, early instar pandora larvae overwinter, then feed to maturity in 
the spring, pupate during the summer and emerge later that same 
summer, or as is often the case, the following summer. Adult velda 
emerge between 0930 and 1100 h. Females begin emitting pheromones 
within one hour after it becomes dark; some females fly prior to mat- 
ing. The developmental pattern of velda explains its early flight and 
suggests how it can occur sympatrically with lindseyi in the San Ber- 
nardino Mts. The main flight of velda occurs about two months prior 
to that of lindseyi. 

Larvae of lindseyi and velda are easily separated. Mature velda 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 










. burnsi 
. eglanterina 
. eglanterina annulata 











. eglanterina shastaensis er 
- Nevadensis » nuttalli 
H. nuttalli uniformis 


H. juno 




















CALIFORNIA INSECT SURVEY 
i ‘ 


Department of Entomology and Parasitology 





CALIFORNIA INSECT SURVEY Give ge 
Department of Entomology and Parasitology y 


UNIVERSITY OF CALIFORNIA UNIVERSITY OF CALIFORNIA 























Fic. 5. The distribution of eight species and four subspecies of Hemileuca in Cali- 
fornia. 


larvae (Figs. 4a, b) are light gray to gray-purple; the dorsal scoli are 
of the rosette type, orange in coloration, and similar in structure to 
those of Hemileuca. The larvae of lindseyi (Figs. 4d, e) have a dark 
brown ground color; the dorsal scoli are brown, and are a simple 
branched type. The larvae of velda are described here for the first 
time. The description is based on material reared from ova which were 
given to the author by Steve McElfresh. The ova were secured from a 
female collected at the type locality, Coxey Meadow, San Bernardino 
Mts., San Bernardino Co., on 25 June 1982 (Figs. 4a, b, c). 


Description of Larval Instars 


FIRST INSTAR. Head: Brown with sparse light brown secondary setae; diameter 1.5 
mm. Body: Length 7.4 mm, width 2.1 mm. Ground color brown. Ventral surface yellow. 
Lateral surface with 3 lateral white bands. Line I encompasses the lateral scoli and 
extends length of larva. Line II, just ventral of dorsal lateral scoli and extends length of 
larva. Lateral surface between band 1 and 2 appears as dark brown band. Line III extends 
length of larva on dorsal side of dorsal lateral scoli. Abdominal dorsal and mid dorsal 
area yellow with remnants of black mid dorsal line. Dorsal thoracic segments dark brown 
to black. Lateral and sublateral scoli with short black spine. Dorsal lateral scoli with 
simple black spine twice length of lateral spine. Dorsal spines on abdominal scoli 1-8 
black with brown setae at tip. Dorsal thoracic and 9th abdominal mid dorsal scoli branched 
with short brown setae extending from tips. Anal shield black, true legs dark brown, 
prolegs yellow. 

SECOND INSTAR: Head: Brown with sparce light brown secondary setae; diameter 2.1 
mm. Body: Length 10.2 mm, width 2.8 mm. Ground color brown. Ventral surface light 
brown. Lateral surface with 3 lateral white lines as in first instar. Brown line connects 
all dorsal scoli. Mid dorsal scoli yellow with thin brown mid dorsal line. Dorsal thoracic 


VOLUME 388, NUMBER 4 307 





Fic. 6. Habitat of desert, sagebrush and coastal chaparral species of Hemileuca in 
California. a, Coastal Chaparral, San Diego Co. Habitat of H. electra electra, Saturnia 
walterorum and Hyalophora eurylaus. Prominent vegetation in photo includes Artemisia 
californica, Adenostoma fasciculatum, Eriogonum fasciculatum, Rhus laurina, and Sal- 
via apiana; b, Great Basin, Mono Co. Habitat of H. eglanterina annulata, H. nuttalli 
uniformis, H. hera hera and Hyalophora gloveri. Prominent vegetation in photo includes 
Artemisia tridentata, Purshia tridentata and Symphoricarpos vaccinioides; e. High Des- 
ert, San Bernardino Co. Habitat of H. electra clio and H. burnsi. Prominent vegetation 
in photo includes Larrea divaricata, Yucca brevifolia, Tetradymia axillaris, Eriogonum 
fasciculatum var. polifolium, and Opuntia bigelovii; d, High Desert Wash, San Bernar- 
dino Co. Habitat of H. neumoegeni, H. electra clio, and Sphingicampa hubbardi. Prom- 
inent vegetation in photo includes Rhus trilobata anisophylla, Prunus fasciculata, Acacia 
greggii, Eriogonum fasciculatum var. poliofolium and Prosopis juliflora. 


area dark brown. Intersegmental areas white. Ventral, lateral and dorsal lateral scoli 
black. Dorsal scoli black on a yellow light brown pedestal. Dorsal, dorsal lateral, and mid 
dorsal scoli branched; thoracic scoli enlarged. True legs black. Anal shield dark brown. 
Prolegs light brown. Spiracles dark brown. 

THIRD INSTAR. Head: Light brown with short white setae; diameter 2.4 to 2.7 mm. 
Body: Length 16 to 17 mm, width 4 mm. Ground color brown. Ventral surface light 
brown. Lateral surface with prominent white undulating line passing through base of 
each sublateral scolus and extending length of larva. Spiracular area light brown. 2nd 
white line extending length of larva, passing just ventral of lateral scoli. Brown line 
extending length of larva, passing through lateral scoli. 3rd white line extending length 
of larva, passing midway between dorsal and lateral scoli. Dorsal area white with thin 
black mid dorsal line. Thoracic and caudal scoli enlarged and branched. Dorsal and 
dorsal lateral scoli black and branched and on yellow pedestal. Sublateral and ventral 
scoli black and unbranched. Thoracic shield brown. Anal shield brown with white stripes. 
True legs black. Prolegs light brown. Spiracles brown. 


308 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


FOURTH INSTAR. Head: Brown with light brown setae, ocellar scar dark brown; di- 
ameter 3.4-3.6 mm. Body: Length 28 to 32 mm, width 6 mm. Ground color gray. Ventral 
surface gray to yellow. Lateral surface with 3 light gray lines. Line I, undulating line 
extending length of larva and passing through base of each sublateral scolus. Line II, 
extending length of larva, passing just ventral of lateral scoli, and diffuse or absent. Line 
III, straight line passing midway between dorsal and dorsal lateral scoli. Mid dorsal area 
with thin dark brown mid dorsal line surrounded on each side by light gray line. Lateral 
and dorsal segmental area between light gray lines dark gray-brown with few light gray 
pinacula. A prominent gray dot occurs just dorsal of lateral scoli on abdominal segment 
1 and occasionally on abdominal segment 7. All scoli branched. Dorsal and dorsal lateral 
scoli on yellow pedestal, base of spines yellow, distal portion black, except for black 
caudal scoli. Thoracic shield brown, anal shield brown with white stripes. True legs dark 
brown. Prolegs light brown to yellow. Spiracles light brown to yellow. 

FIFTH INSTAR. Head: Light brown with few short white or hylen setae. Ocellar scar 
black. Clypeus brown. Mandibles dark brown. Diameter 4.5 to 5.5 mm. Body: Length 
49 to 57 mm, width 10-12 mm. Ground color gray to gray brown. Ventral, lateral, and 
dorsal surface gray. Undulating subspiracular light gray fold extending length of larva. 
Lateral and dorsal surface with numerous folds on segmental and intersegmental area; 
brown line occurring at bottom of each fold. Dorsal and lateral surface sparsely covered 
with short hylen secondary setae. Diffuse light orange-brown area passing along base of 
dorsal scoli, extending length of larva, and surrounding light gray mid dorsal line. Shaft 
of all scoli orange. Sublateral, lateral and dorsal lateral scoli branched with white and 
dark brown spines. Dorsal scoli rosette type with white spines and black tips. Thoracic 
and anal shields light brown. True legs dark brown. Prolegs gray. Spiracles orange. 


ACKNOWLEDGMENTS 


I would like to thank the following individuals who responded to my requests for 
distributional and flight period data from their personal collections: Mike Collins, Ken 
Hansen, John Johnson, Sterling Mattoon, Steve McElfresh, Scott Meredith, and Mike 
Smith. Special thanks to Ken Hansen, Scott Meredith, Mike Smith and Steve McElfresh 
for providing material any time it was requested and to Steve McElfresh and Jim Tuttle 
for reviewing the manuscript. 


LITERATURE CITED 


AINSLIE, C. N. 1910. The New Mexico Range Caterpillar. U.S. Dept. Agric. Bureau 
Ent. Bull. 85(5):59-96. 

ALDRICH, J. M. 1911. Larvae of saturniid moth used as food by California Indians. J. 
N.Y. Entomol. Soc. 20:28-31. 

1921. Coloradia pandora Blake, the moth of which the caterpillar is used as 
food by Mono Lake Indians. Ann. Entomol. Soc. Amer. 14:36-38. 

CAROLIN, V. M. 1971. Extended diapause in Coloradia pandora Blake (Lepidoptera: 
Saturniidae). Pan-Pacific Entomol. 47:19-23. 

CHAMBERLIN, W. J. 1922. A new lepidopterous enemy of yellow pine in Oregon. J. 
N.Y. Entomol. Soc. 30:69-71. 

COLEMAN, J. 1983. A thousand Hemileuca in one night! Lepid. Soc. News (2):36-37. 

COLLINS, M. M. & P. M. TuskEs. 1979. Reproductive isolation in sympatric species of 
dayflying moths (Hemileuca: Saturniidae). Evolution 33(2):728-733. 

COMSTOCK, J. A. & C. M. DAMMERS. 1937. Notes on the early stages of three California 
moths. Bull. So. California Acad. Sci. 36:68-78. 

1939. Studies in the metamorphoses of six California moths. Bull. So. California 
Acad. Sci. 37:105-128. 

FERGUSON, D. C. 1971. The moths of America north of Mexico. Fascicle 20.2a Bom- 
bycoidea (in part). Classey, London. Pp. 1-154. 

1972. The moths of America north of Mexico. Fascicle 20.2b Bombycoidea (in 

part). Classey, London. Pp. 155-275. 











VOLUME 38, NUMBER 4 309 


HUDDLESTON. E. W., E. M. DRESSEL & J. G. WaTTsS. 1976. Economic threshold for 
range caterpillar larvae on blue grama pasture in northeastern Lincoln County, New 
Mexico, in 1975. New Mexico State Agri. Exper. Station. Research Report 314. 

JOHNSON, J. W. & E. WALTER. 1980. A new species of Coloradia in California (Satur- 
niidae, Hemileucinae). J. Res. Lepid. 18(1):60-66. 

MCFARLAND, N. 1974. Notes on three species of Hemileuca from eastern Oregon and 
California. J. Lepid. Soc. 28(2):136-141. 

MICHENER, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemisphere, 
morphology, phylogeny, and classification. Bull. Amer. Mus. Nat. Hist. 98(5):335- 
502. 

PACKARD, A. S. 1914. Monograph of the bombycine moths of North America, part 38. 
Mem. Nat. Acad. Sci. 12. Pp. i-ix, 1-276, 503-516. 

PATTERSON, J. E. 1929. The Pandora moth, a periodic pest of western pine forests. 
U.S.D.A. Tech. Bull. 187. 19 pp. 

SCHAAF, R. 1980. Forest Service team to assess Pandora Moth effects. United States 
Forest Service, Inyo National Forest News, 11-4-80. 

1981. Pandora Moth outbreak in sharp decline. U.S. Forest Service, Inyo Na- 
tional Forest News, 4-29-81. 

SMITH, M. J. 1974. Life history notes on some Hemileuca species (Saturniidae). J. Lepid. 
Soc. 28(2):142-145. 

TusKES, P. M. 1976. A key to the last instar larvae of west coast Saturniidae. J. Lepid. 
Soc. 30(4):272—276. 

1978. A new species of Hemileuca from the southwestern United States (Sa- 
turniidae). J. Lepid. Soc. 32(2):97-102. 

Watts, J. G. & T. D. EVERETT. 1976. Biology and behavior of the range caterpillar. 
New Mexico State Agri. Exper. Station Bull. 646. 

WyGANT, N. D. 1941. An infestation of the Pandora Moth, Coloradia pandora Blake, 
in lodgepole pine in Colorado. J. Econ. Entomol. 34(5):697-702. 








Journal of the Lepidopterists’ Society 
38(4), 1984, 310-316 


OBSERVATIONS ON THE BIONOMICS OF 
HELIOTHIS PHYLOXIPHAGA (NOCTUIDAE) ON CLUSTER 
TARWEED IN SOUTHEASTERN WASHINGTON! 


G. L. PIPER AND B. L. MULFORD? 


Department of Entomology, Washington State University, 
Pullman, Washington 99164 


ABSTRACT. The bionomics and brief descriptions of the life stages of Heliothis 
phyloxiphaga Grote & Robinson, a noctuid associate of the composite weed, Madia 
glomerata Hook., are presented. The moth was univoltine in southeastern Washington, 
with peak adult populations appearing during late July. Eggs were deposited on imma- 
ture involucres, and the first- and second-stage larvae fed within the involucres on the 
developing achenes, while later stage larvae consumed bracts, flowers, and leaves. Pu- 
pation occurred in the soil from late August to early October. Under laboratory condi- 
tions, the life cycle of the moth from egg to adult required ca. 52 days. 


The entomofaunas of many plants indigenous to North America are 
either unknown or inadequately characterized. Typically, little in-depth 
biological information is available on specific associates. This is partic- 
ularly true for cluster or stinking tarweed, Madia glomerata Hook. 
(Compositae: Madiinae), a weed of the western United States and Can- 
ada. 

The plant is an erect, 30-100 cm tall, yellow-flowered, summer an- 
nual that often forms dense stands in rangeland, pastures, along road- 
sides or ruderal areas (Dennis, 1980). Numerous stalked glands on the 
stems, leaves, and floral capitula produce a nauseating, tar-scented, 
sticky exudate that is readily transferred to clothing or animals upon 
contact. M. glomerata is rarely grazed by livestock and thus has vir- 
tually no forage value. The weed is allelopathic and prevents estab- 
lishment or reduces growth of various desirable forbs and grasses (Car- 
nahan & Hull, 1962). 

In southeastern Washington, cluster tarweed was consistently at- 
tacked by the noctuid Heliothis phyloxiphaga Grote & Robinson. Al- 
though much biological information is available for the more impor- 
tant agricultural pest species of Heliothis such as H. virescens (F.) and 
H. zea (Boddie) (Hardwick, 1965), very little is known about the bio- 
nomics of the non-economically important species including H. phy- 
loxiphaga. This lack of information prompted the present investiga- 
tion. 


‘Scientific paper no. 6727. Work conducted under projects 0335 and 0582, Washington State University, Agricultural 
Research Center, Pullman. 
* Current address: Boyce Thompson Southwestern Arboretum, P.O. Box AB, Superior, Arizona 85273. 


VOLUME 38, NUMBER 4 311 


METHODS 


Studies on the biology of H. phyloxiphaga were conducted in the 
laboratory and correlated with observations made in the field at several 
sites within a 15 km radius of Pullman (Whitman County) in south- 
eastern Washington from 1979-83. Laboratory rearings, initiated from 
field-collected eggs and larvae, were maintained at 20-25°C, 40-50% 
RH, and 14 L:10 D hour regime. Eggs were confined to 10.0 x 1.5 
cm plastic petri dishes lined with moistened paper towelling until 
hatching occurred. Newly emerged to mid-fifth stage larvae were reared 
individually in petri dishes provisioned with a moistened paper sub- 
strate and sufficient host plant material. The paper and food were 
replaced daily. Nearly mature larvae were transferred to ventilated 
7.0 x 8.5 X 3.5 cm clear plastic boxes filled with sandy loam soil for 
pupation. Pupae remained in the cages until adult eclosion. 

Emergent moths were sexed, paired, and placed in cylindrical (15 
cm diam x 80 cm high) translucent plastic cages ventilated apically 
with fine mesh saran screen. A 2 dr shell vial filled with a 20% honey- 
water solution and plugged with cotton dental wicking along with 
several freshly-excised, field-collected M. glomerata floral shoots in- 
serted through the Parafilm® seal of a water-filled receptacle were 
placed in the cage and replenished as needed. Nocturnal observation 
of adults was accomplished under red illumination (simulated dark- 
ness). 


RESULTS 


Distribution. H. phyloxiphaga has a distribution that ranges from 
the Mississippi River west to California, north to the Canadian prov- 
inces of Alberta and British Columbia, and south to Mexico, with ad- 
ditional records from Illinois, Massachusetts, and New York (Forbes, 
1954; Crumb, 1956). 

Host plants. Based upon an examination of pertinent literature, the 
moth has a broad host range. In addition to M. glomerata, the follow- 
ing larval food plants have been noted by Grote and Robinson (1867), 
Crumb (1926, 1956), Forbes (1954), Tietz (1972), and T. F. Watson 
(pers. comm., Univ. Arizona): COMPOSITAE: Achillea millefolium 
L., Balsamorhiza sp., Chaenactis douglasii (Hook.) H. & A., Erigeron 
divergens T. & G., Grindelia camporum Greene, G. robusta Nutt., G. 
squarrosa (Pursh) Dunal, Hemizonia congesta DC., Lactuca sativa L., 
Machaeranthera canescens (Pursh) Gray, Parthenium argentatum 
Gray; GERANIACEAE: Erodium cicutarium (L.); GRAMINEAE: 
grasses; IRIDACEAE: Gladiolus sp.; LEGUMINOSAE: Lathyrus sp., 


312 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Medicago sativa L.; POLEMONIACEAE: Gilia aggregata Spreng., 
Phlox sp.; ROSACEAE: Fragaria sp.; SCROPHULARIACEAE: Antir- 
rhinum sp.; and SOLANACEAE: Schizanthus sp. 

Description of stages. Egg. The pearly pale yellow hemispherical 
egg averaged 0.63 + 0.25 mm in height and 0.69 + 0.03 mm in 
diameter (n = 25). The chorion was sculptured with numerous prom- 
inent ribs that radiated from a nipple-like micropyle positioned at the 
apical pole. A reddish purple band developed in the micropylar half 
of the egg within 72 h of deposition, and as embryogenesis proceeded 
the entire egg assumed a brownish grey color. 

Larva. The first-stage larva was creamy grey with a blackish brown 
head capsule and prothoracic shield. The second-stage larva differed 
little except for a somewhat paler head capsule, prothoracic shield, and 
greenish grey color. Definitive maculation and coloring become evi- 
dent in the third-stage larva and are maintained in fourth- and fifth- 
stage larvae. The fifth-stage larva had a greenish brown to beige head 
capsule mottled with brown and a light green or olive-green body. An 
excellent description of the mature larva is given by Crumb (1926, 
1956) and abbreviated descriptions occur in Lange and Michelbacher 
(1937) and Stahler (1939). Lange and Michelbacher (1937) also includ- 
ed a photograph of the mature larva and pictured the larval chaeto- 
taxy. 

The body lengths of ten first- through fifth-stage larvae averaged 
2.72 + 0.48 mm, 5.62 + 0.75 mm, 12.09 + 3.89 mm, 19.56 + 1.29 
mm, and 28.15 + 3.30 mm, respectively. First- and second-stage larvae 
experienced an average increase in length of 47% between successive 
molts, whereas, third- to fifth-stage larvae grew 65%. Mean cast head 
capsule widths of the five instars were 0.32 + 0.02 mm, 0.52 + 0.05 
mm, 0.90 + 0.10 mm, 1.61 + 0.06 mm, and 2.55 + 0.16 mm (n = 
15), respectively. 

Pupa. The pupa was obtect and glossy golden amber when first 
formed but became chestnut-brown as it matured. In the male, the 
slit-like genital opening bordered by a pair of tubercles was situated 
mid-ventrally on the ninth abdominal segment. In the female, the 
genital aperture occurred on the eighth abdominal segment. The cre- 
master consisted of two slightly curved, divergent spines arising from 
the conical apex of the tenth abdominal segment. Lange and Michel- 
bacher (1937) provided a photograph of the pupa. The mean length 
of 17 pupae was 16.89 + 3.96 mm. 

Adult. Adult H. phyloxiphaga males and females are similar in ap- 
pearance. When viewed from above, the body and forewings are a 
dull brownish yellow with a distinctive olive-green tinge, the forewings 
being marked with several brownish black spots and brown to greenish 


VOLUME 38, NUMBER 4 3138 


fawn irregular transverse bands. The maculation of the creamy buff 
hindwings consisted of a dull black discal spot and a black, broad, 
transverse marginal band interrupted by a central creamy buff patch. 
Detailed descriptions of the adult are given by Grote and Robinson 
(1867) and Forbes (1954). Lange and Michelbacher (1937) included a 
photograph of a series of adults showing subtle color and maculation 
variations and also illustrated the male genitalia. The mean body length 
and alar expanse of 25 adults was 14.5 + 0.6 mm and 33.0 + 1.0 mm, 
respectively. 

Life history and habits. In the southernmost portions of its range, 
H. phyloxiphaga may be bivoltine (Lange & Michelbacher, 1937; 
Forbes, 1954), with first and second generation adults appearing from 
April-May and July—August, respectively. In Washington the moth was 
univoltine. Based upon field observations and examination of label data 
from specimens deposited in the M. T. James Entomology Collection, 
Washington State University, adult activity commenced in late June 
and extended to early September, with peak populations being record- 
ed from mid-July to early August. Crumb (1956) reported similar find- 
ings regarding seasonal occurrence of the adults. 

Adults were not commonly encountered in nature. The moths are 
inactive during the day and were occasionally observed clinging to 
stems of dead weeds or grasses and amongst leaf litter, the coloration 
of the closed forewings blending imperceptibly with the resting sub- 
strate, presumably affording the adult protection from predation. If 
disturbed, adults flew short distances before resettling. Adult feeding 
was not observed in the field, but individuals readily imbibed the 
honey-water provided in the laboratory. 

Mean longevity of 24 laboratory-reared males was 13.7 + 6.8 (range 
5-30) days; longevity of females was 12.5 + 8.9 (range 4-29) days. 
The sex ratio of 44 cultured adults was 1.2:1 (24 males:20 females). 

Mating behavior was not observed for H. phyloxiphaga but it is 
probably similar to that described for H. virescens (Lingren et al., 
1977) and H. zea (Callahan, 1958). 

The preoviposition period, from emergence to first deposition of 
eggs, for six females averaged 6.2 + 2.8 (range 4-8) days. Based only 
upon days when these females oviposited, they laid an average of 
29.8 + 26.4 (range 1-130) eggs per day over a 12.0 + 6.9 (range 4- 
23) day period, with the majority of eggs being deposited during the 
first seven days. Total eggs laid per female ranged from 53-536, the 
average being 282.3 + 190.6. 

Oviposition was generally a crespuscular activity. Plants averaging 
40.0 + 10.8 (range 17.5-62.5) cm in height (n = 125) with terminally 
developing inflorescences were selected for oviposition purposes. When 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


an acceptable ovipositional site was encountered by a mated female 
she momentarily alighted on a cluster of flower heads and rapidly 
vibrated her wings prior to egg deposition. Wing movement ceased as 
the female appressed the tip of her abdomen to the outer surface of 
an involucral or receptacular bract to which she affixed an egg, the 
deposition process requiring only one or two sec. The female resumed 
flight and laid additional eggs on the same plant or on nearby plants. 
In the laboratory, females would often feed and/or rest for brief in- 
tervals during the oviposition period. Field observations revealed that 
females deposited nearly equal numbers of eggs on the bracts of the 
involucre or receptacle. Of 28 eggs laid on involucres, 16 (57%) were 
attached to the central head in a cluster, the others being affixed to 
peripheral heads. Egg distribution on 100 field-examined plants was as 
follows: 60 had a single egg, 24 had 2 eggs, nine had 3 eggs, four had 
4 eggs, five had 2 eggs, and one plant had 6 eggs. Eggs were found in 
nature from mid-July to the first week in September. 

Prior to hatching, the larva assumed a U-shape within the eggshell, 
with the head and caudal end tightly compressed at the micropylar 
end. The larva chewed a ragged hole through the chorion in the polar 
or equatorial region. When the orifice approximated the size of the 
head capsule, the larva exited by peristaltic contractions of its body. 
The entire hatching process required ca. 12-15 min. The chorionic 
remnant was generally not consumed by the neonatal larva. The in- 
cubation period of 48 laboratory-laid eggs was 5.4 + 0.5 days. Viability 
of these eggs exceeded 98%. 

A newly emerged larva initially fed sparingly upon the epidermis 
and underlying parenchyma cells of the bracts upon which the egg 
had been laid. Soon thereafter the larva crawled to and chewed a hole 
through an involucre near its mid-point or base and entered, wherein, 
it fed upon the developing florets and immature achenes. A sparse, 
irregular deposition of silk was evident upon and amongst the stalked 
glands of the infested involucre. The webbing and extruded frass pel- 
lets provided concealment for the feeding larva. Each cluster tarweed 
involucre contained ca. 12 (range 8-20) achenes that were destroyed 
or damaged by larval feeding. A first- and second-stage larva was each 
capable of destroying four to five involucres. Attacked involucres shri- 
veled and failed to open. Third- to fifth-stage larvae were external 
feeders on flowers, involucral and receptacular bracts, and leaves but 
would not eat achenes approaching maturity. These larvae, especially 
in dense stands of the weed, moved from plant to plant feeding upon 
the succulent tissues and defoliating the flowering shoots. Larvae of 
this age category were highly cannabalistic, and consequently, only a 


VOLUME 38, NUMBER 4 315 


single larva per plant was found in nature. The green larvae were not 
easily discernible among foliage and flower heads, their cryptic col- 
oration probably affording a degree of protection from predators. Old- 
er larvae, when disturbed, either regurgitated a droplet of green fluid 
or dropped from the plant to the soil, assumed a coiled posture, and 
remained motionless for several minutes. 

The duration of the first to fifth stadia averaged 4.0 + 0.3 (range 3- 
5) days (n = 53), 3.7 + 0.7 (range 3-5) days (n = 48), 4.0 + 0.7 (range 
3-6) days (n = 34), 6.2 + 1.0 (range 5-8) days (n = 26), and 7.9 + 1.0 
(range 6-9) days (n = 19), respectively. 

The fifth-stage larva stopped feeding during the sixth to ninth day 
of development. The larva became strongly positively geotactic and | 
descended to the soil and burrowed to a depth of 1.5-4.0 cm. Having 
reached a suitable depth, the larva constructed an emergence tunnel 
and elliptical pupal cell. The smooth-walled, sparsely silk-lined cham- 
ber averaged 20 mm in length and 10 mm in diam (n = 20). The 
upward-sloping emergence tunnel was sealed with a loosely fitting plug 
of silk-bound soil particles. 

Upon completion of the cell, the larva contracted to ca. 50% of its 
former length (x = 18.5 + 2.29 mm; n = 8), assumed a lime green 
color, and became quiescent. About 24 h later, the prepupal integu- 
ment ruptured medially along the dorsum of the thorax and the head 
of the pupa emerged through the slit. The integumental remnant was 
slid posteriorly by body movement of the pupa and eventually formed 
a loose mass at the base of the cremaster. Pupal emergence was com- 
pleted in ca. five min. The duration of the prepupal period, from 
feeding cessation to pupa formation, was 3.2 + 0.4 days (n = 20). 

The pupa was usually positioned within the cell with the head di- 
rected toward the emergence tunnel. If physically disturbed, the pupa 
slowly rotated its abdomen in a clockwise direction. 

In the field, pupation occurred from late August to early October, 
the pupa overwintering in an obligate diapause state. The pupal period 
of 59 laboratory-reared, non-diapausing individuals averaged 17.5 + 
2.1 (range 13-23) days. 

Adult eclosion was facilitated through fractures in the pupal exu- 
vium, which developed dorsally along the thorax and ventrally near 
the antennae. The adult then freed itself by alternate expansions and 
contractions of the abdomen and leverage afforded by the legs. Emer- 
gence took place within the pupal cell, and the crumpled winged adult 
exited through the tunnel to the soil surface where wing expansion and 
integument hardening was accomplished. Adult emergence was a noc- 
turnal event, occurring between 2100 and 0300 h. 


316 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Natural enemies. The anthocorid, Orius minutus (L.), was observed 
feeding on the eggs of H. phyloxiphaga. However, the extent to which 
this predator destroyed eggs was not determined. 

An unidentified braconid, a primary solitary larval parasitoid, was 
responsible for ca. 25% of the late stage larval mortality observed in 
nature. Intraspecific, internecine combat also contributed to larval 
mortality as previously noted. 

No pupal or adult parasitoidism or predation was observed during 
the study. 


ACKNOWLEDGMENTS 


The authors wish to thank R. W. Poole, USDA Systematic Entomology Laboratory, 
IIBIII, for confirmation of our tentative identification of H. phyloxiphaga. We also ex- 
press appreciation to D. Thompson and L. Walls for their invaluable assistance with field 
collections and laboratory rearings. 


LITERATURE CITED 


CALLAHAN, P. S. 1958. Behavior of the imago of the corn earworm Heliothis zea 
(Boddie), with special reference to emergence and reproduction. Ann. Entomol. Soc. 
Am. 51:271-283. 

CARNAHAN, G. & A. C. HULL, Jr. 1962. The inhibition of seeded plants by tarweed. 
Weeds 10:87-90. 

CRUMB, S. E. 1926. The Nearctic budworms of the lepidopterous genus Heliothis. Proc. 
U.S. Nat. Mus. 68:1-8. 

1956. The larvae of the Phalaenidae. USDA Tech. Bull. 1135:1-356. 

DENNIS, L. J. 1980. Gilkey’s Weeds of the Pacific Northwest. Oregon State Univ. Press, 
Corvallis. 382 pp. 

ForBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Part III. 
Cornell Univ. Agric. Expt. Sta. Mem. 329:1-483. 

GrRoTE, A. R. & C. T. ROBINSON. 1867. Descriptions of American Lepidoptera. Trans. 
Am. Entomol. Soc. 1:171-192. 

HARDWICK, D. F. 1965. The corn earworm complex. Mem. Entomol. Soc. Can. 40:1- 
247. 

LANGE, W. H. & A. E. MICHELBACHER. 1937. Two closely related species of Heliothis 
found in tomato fields of central California. Bull. Calif. Dept. Agric. 26:320-325. 

LINGREN, P. D., G. L. GREENE, D. R. Davis, A. H. BAUMHOVER & T. J. HENNEBERRY. 
1977. Nocturnal behavior of four lepidopterous pests that attack tobacco and other 
crops. Ann. Entomol. Soc. Am. 70:161-167. 

STAHLER, N. 1939. Notes on the taxonomy of noctuid larvae. Pan-Pacific Entomol. 15: 
123-126. 

TIETZ, H. M. 1972. Index to the described life histories, early stages and hosts of the 
Macrolepidoptera of the continental U.S. and Canada. Allyn Mus. Entomol., Sarasota, 
FL. Vol. 1:1-536. 





Journal of the Lepidopterists’ Society 
38(4), 1984, 317-318 


GENERAL NOTES 


AN UNUSUAL OVIPOSITIONAL SITE FOR 
AMPHIPYRA TRAGOPOGINIS (L.) (NOCTUIDAE) 


Egg clusters (4-12 eggs) of Amphipyra tragopoginis (L.) were found within the cavities 
of 30 helicoid cases of Apterona helix (Siebold) (Lepidoptera: Psychidae) (Fig. 1) during 
observations in Lenox, Massachusetts of the presence of a large aggregation of these 
garden bagworm larval cases attached to above ground substrates including: domiciles, 
wood fencing, grasses, weeds, shade trees, ornamentals, planted flowers and vegetables. 

Although samples of the helicoid cases were collected from the above mentioned 
substrates, the only cases containing eggs were collected from untreated wood fencing. 

These observations suggest that tiny crevices on tree trunks and branches or “artificial” 
above ground surfaces may be used by gravid A. tragopoginis females as oviposition 
sites. Two closely related species, Amphipyra pyramidoides (Guen.) and A. glabella 
(Morr.), oviposit in crevices on trees and twigs of their hosts (J. G. Franclemont, pers. 
comm.), indicating that members of this genus prefer similar oviposition sites. 

It is not suggested that A. helix is an “obligate” in the life cycle of A. tragopoginis 
but that the cases by chance provide an appropriate stimulus for oviposition behavior by 
A. tragopoginis. 

The identity of A. tragopoginis was confirmed by using adults obtained from larvae 
reared on artificial diet. 





Fic. 1. SEM of egg cluster of Amphipyra tragopoginis (L.) (note arrow) within cavity 
of larval case of Apterona helix (Siebold). 33x. Scale line = 1 mm. 


318 aie JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


D. ADAMsKI, Department of Entomology, University of Massachusetts, Amherst, 
Massachusetts 01003. Present address: Department of Entomology, Mississippi State 
University, Mississippi State, Mississippi 39762. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 318-319 


ACER NEGUNDO (BOXELDER) AS A FOOD PLANT FOR 
SYNANTHEDON ACERRUBRI (SESIIDAE) 


On 30 June 1983 quite by accident I discovered a Synanthedon acerrubri (Engelhardt) 
ovipositing on a somewhat distressed Acer negundo (boxelder) (Aceraceae) in my yard 
in Liberty, Missouri. A total of 18 acerrubri were caught through the 16th of July 1983. 
At least one specimen was caught on all of the intervening days except the 8th and 15th, 
two days on which I did no collecting at all. Eight females and ten males were caught. 
Prior to this only seven males had been collected in Missouri, to the best of my knowledge. 
All of my specimens were caught on one of three A. negundo growing in my yard. They 
favored the most distressed tree. Other trees in the yard and neighborhood were checked 
for visitation. This included other members of the maple family, plus cherry, pear, elm, 
hackberry and cottonwood. 

Initially, I tried to catch the specimens with a net on which I had pinned male sex 
attractant. No males seemed attracted to the bait at that time or when I subsequently 
pinned the attractant to my shirt during my collecting. Almost all of the specimens were 
caught with a small killing jar, the moths being taken directly off the trees. 

Most were seen ovipositing or resting no more than a foot from the ground on the 
bark of the host tree. Only three were seen or caught at a height above 4 ft. All except 
one specimen were caught between 1430 and 1900 h. They appeared most commonly 
around 1730 h. None was seen mating. A number of pupal cases were found projecting 
from the trunk of the most distressed looking A. negundo and appeared similar to that 
pictured in Holland (1903, The moth book, Doubleday, Page & Co.) for Synanthedon 
acerni (Clemens). It would seem logical to assume that these were pupal cases of the S. 
acerrubri, but none was seen emerging. 

This may not be the normal time of emergence since it was a very late year for many 
Lepidoptera species in Missouri. J. Richard Heitzman (pers. comm.), who collected the 
other recorded specimens in Missouri of which I am aware, captured all of them in his 
yard in Independence. He caught five male acerrubri nectaring at Asclepias syrica (pur- 
ple milkweed, Asclepiaceae) between the 10th and 29th of June over a number of years, 
and two male specimens were collected while responding to a specific sex attractant at 
approximately 1145 h, 11 July 1982 and 9 July 1983. These latter dates correspond nicely 
with the dates that I collected the 18 specimens. 

In 1984 a total of 63 acerrubri were caught at the two most distressed boxelder in my 
yard. All were caught between 1600 and 2020 h from 9 June to 18 July. Three males 
came to an attractant at a different location in the yard. They were caught while resting 
on leaves at this location (one at 1600 h on 2 July and two at 1820 h on 9 July). A total 
of 65 acerrubri (31 males and 34 females) were caught. One specimen was sighted but 
not caught on 24 July. Once again pupal cases (16) were seen projecting from the trunks 
of the A. negundo. 

S. acerrubri occurs in the eastern United States and is known to feed on Acer rubrum 
(red maple) and A. saccharum (sugar maple), according to Engelhardt (1946, The North 
American clear-wing moths of the Family Aegeriidae, U.S. Nat. Mus. Bull. 190). A. 
negundo is apparently an unrecorded food plant for acerrubri. Since the growth of 


VOLUME 88, NUMBER 4 319 


boxelder is widespread, the moth may be more common than previously thought but has 
gone undetected. 

I am grateful to J. Richard Heitzman for his aid in identifying and determining the 
sex of the specimens and for being so generous with his time and knowledge. My son, 
James Adams, deserves my thanks for reviewing this manuscript. 


ELEANER R. ADAMS, Biology Department, William Jewell College, Liberty, Missouri 
64068. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 319-322 


ON THE ORIGIN OF SNOUT BUTTERFLIES 
(LIBYTHEANA BACHMANII LARVATA, LIBYTHEIDAE) 
IN A 1978 MIGRATION IN SOUTHERN TEXAS 


Southern Texas periodically is the scene of migrations by the snout butterfly, Liby- 
theana bachmanii larvata (Strecker). The last massive migration in Texas occurred dur- 
ing summer 1971 (Helfert, 1972, Entomol. News 83:49-52; Neck, 1983, J. Lepid. Soc. 
37:121-128). More frequent than these “cloud-type” migrations are the smaller-scale 
migrations which rarely extend beyond the northeastern boundary of the South Texas 
Plains (line from San Antonio to the Gulf Coast north of Corpus Christi). A series of 
these more restricted migrations was observed during four traverses of the area in June, 
July and September of 1978. Comments from two observers will be integrated into 
personal observations. The primary thrust of the investigation of this migration was to 
determine the geographical origin of the migrating butterflies. A secondary thrust was 
to document a relationship between density of butterfly flights and local habitat. 

28 June 1978. On the Coastal Plain of Texas, a low-density migration was observed 
from north of Refugio, Refugio Co., to south of Sinton, San Patricio Co. (Fig. 1). Density 
of migratory snout butterflies varied with vegetation and urban/rural settings (Table 1). 
Snout butterflies were not very common over recently-harvested sorghum fields, were 
most abundant in areas of invaded brush patches (dominated by mesquite, Prosopis 
glandulosa), and were less common but not absent from urban areas, e.g. Woodsboro (a 
small farming community center). Note should be made that some brush plots exhibited 
no flying butterflies. 

Most snout butterflies were flying an approximate west-to-east flight path. Azimuth 
directions of compass heading of butterflies at four localities were as follows: 1) Refugio, 
110°; 2) Sinton, 115°; 3) IH387/US 77 bridge over Nueces River, 85°; and 4) 5 km south 
of Kingsville, 80°. These flight lines were extended inland in an attempt to discover 
source regions of these butterflies. While no information is available on the distance flown 
by these butterflies, flight lines (Fig. 1) indicate a broad source area for the observed 
snout butterflies. Several butterflies were observed moving westward temporarily as a 
result of vehicle-caused air turbulence. Several butterflies were observed being forced 
along the axis of the highways when two semi-trailer trucks approached and passed each 
other. The only other butterfly species associated with the migrating snouts were occa- 
sional specimens of the queen, Danaus gilippus strigosus (Bates), which totaled less than 
five percent of the total butterfly count. 

2 July. Traveling northward from Brownsville, Cameron Co., the first snout butterflies 
were encountered just north of Kingsville. Butterflies were traveling eastward (exact 
azimuths not measured) and were common to Robstown and Mathis. Snout butterflies 


320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


e PEARSALL 


ROBSTOWNe 


KINGSVILLE, _» 


PREMONT 


I 


{ LINN-SAN MANUEL 
e@ 


S 
~ 
Ww 
= 
S 
w 
S 
S 


MEXICO 
EDINBURG 


Grande 
20 30 MILES 


32 48KiILOMETERS 





Fic. 1. Map of southern Texas with localities of observations and flight directions of 
Libytheana bachmanii larvata. 


were extremely abundant southeast and north of Oakville. Flight directions were vari- 
able, but the majority were flying somewhat south of east (Fig. 1). A sample of the snout 
butterflies revealed that freshly-emerged adults of both sexes were migrating (no sex- 
ratio recorded). Associated species were (Kricogonia lyside (Godart) (yellow and white 
morph), Eurema lisa Boisduval and Le Conte, and Phoebis sennae (Linnaeus). These 
species were much less common than the snouts, which represented approximately 99% 
of the migrant butterflies. 


VOLUME 38, NUMBER 4 321 


TABLE 1. Number of snout butterflies, Libytheana bachmanii larvata, observed in 
highway driving counts, summer 1978. 


Highway length 
(km) 


Habitat Butterfly number 
28 June 
Woodsboro 1.1 5 
Nueces River bridge 0.5 8 
Cut sorghum field 0.5 0 
Brush plot 0.5 15 
26 September (south of George West)—paired habitats 
Pasture-Brush 0.6 each 5-9 
Pasture-Brush 0.6 each 12-24 
Pasture-Brush 0.6 each 39-47 


Observation by informants. A letter dated 28 June 1978 from George Toalson of 
Pearsall, Frio Co., reported “thousands” of snout butterflies which exhibited no “partic- 
ular direction in their flight pattern” and were observed to “congregate at damp places 
on roadsides and on anything that has flowers on it.’ Phone conversation with Toalson 
revealed a slow buildup in the Pearsall area. Rains had occurred in mid-May and early 
June. Common plants visited were virgin’s bower (Clematis drummondii) and cowpen 
daisy (Verbesina encelioides). Many snout butterflies had been killed by pesticide which 
had been sprayed on a field of peas; the “ground appears brown with bodies.” Note that 
snout butterflies had been observed in 1971 in Austin, Travis Co., feeding at internodes 
of bean plants (Neck, op. cit.). 

J. Stephen Neck reported to me on 10 July that he observed no snout butterflies 
between Brownsville and Austin (via highways 77, 181, 80 and 188) on that day. 

26 September. Driving south from Austin, snout butterflies were first encountered 
near Oakville where movements were in varied directions. In the region from 8 to 18 
km south of George West, Live Oak Co., snout butterflies generally were traveling in 
directions between 105° and 125°. At a point 29 km south of George West (Live Oak- 
Jim Wells County line), they were flying as high as 8 m above the soil surface, although 
some individuals were observed to land upon the road bed (U.S. 281). Also present were 
a number of Phoebis sennae, some of which exhibited courtship behavior. 

A series of paired brush and pasture segments between 80 and 85 km south of George 
West in Jim Wells Co. revealed greater numbers of snouts crossing the highway in areas 
with native brush communities than areas with pastures (Table 1). The effect, however, 
appears to lessen as butterfly density increases. In this area brush communities are dom- 
inated by brasil (Condalia hookeri), whitebrush (Aloysia gratissima), mesquite, retama 
(Parkinsonia aculeata), guayacan (Porlieria angustifolia) and granjeno (Celtis pallida). 
This last species is the prime larval foodplant of the snout butterfly in southern Texas. 
Pastures are dominated by buffelgrass (Cenchrus ciliaris), an exotic grass native to Africa. 
Butterflies flying at the same sites included Euptoieta claudia (Cramer), Eurema lisa, 
Zerene cesonia (Stoll), Phoebis sennae, Papilio cresphontes Cramer, Danaus gilippus 
and Anaea andria Scudder. Also present were substantial numbers of the green darner 
dragonfly, Anax junius. A few snout butterflies were observed chasing D. gilippus. 

Moving southward I approached the southeastward edge of the area containing large 
numbers of snout butterflies. A few were seen in Premont, but none were seen in Fal- 
furrias. West of Falfurrias on Farm-to-Market 285, snout butterflies were abundant. Most 
individuals were moving approximately southward, although some were flying in the 
opposite direction. One individual altered its flight direction by 180° when approached 
by a group of 10 to 12 snout butterflies. Migrating snout butterflies had been observed 


322 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


by local residents for three days. South of Falfurrias in the vicinity of Rachal, they were 
much less common. Very few were observed at Linn-San Manuel. No snout butterflies 
were observed in the Lower Valley between Edinburg and Brownsville. 

29 September. Traveling northward from Brownsville again, no snout butterflies were 
observed until an area north of Mathis (on Farm-to-Market 359) was reached. In the 
area south of Mathis, a large number of butterflies were observed crossing the highway; 
over 90% were Danaus gilippus. Anax junius was once again common. Even in the 
Mathis area, snout butterflies were not common. 

Origin of migrating snout butterflies. Observation of flight directions of snout butter- 
flies in areas with near unidirectional movements (Fig. 1) indicates that some area of 
inland southern Texas was the source area of the 1978 snout migration. Observations 
along the margins of this inland area revealed large scale movements without concen- 
trated peaks of compass directions, but with a tendency for flight away from an area 
still further inland. Backward tracing of flight directions and the observations of Toalson 
indicates a source area south of San Antonio which includes the vicinity of Pearsall. Brush 
communities in this area of southern Texas contain large percentages of Celtis pallida, 
the favored larval foodplant of the snout butterfly. Low densities of snout butterflies 
observed south of Falfurrias are indicative of the lack of C. pallida in the Llano Mesteno, 
a large area of mixed grassland and savannah. 

Analysis of previous snout butterfly migrations (e.g. Neck, op. cit.) has revealed an 
association with exceptionally heavy rainfalls. A large, cloud-type migration in 1971 
followed widespread, heavy rains in southern and central Texas. These summer rains 
followed the intense drought of 1970 and 1971. 

The 1978 migration discussed above is believed to be related to precipitation patterns, 
although the system operating in 1978 differed significantly from 1971. The period from 
summer 1977 to spring 1978 was characterized by rainfall deficiencies (Climatological 
data—Texas, U.S. Department of Commerce). Drought conditions were not as severe as 
the situation present in early summer 1971, however. Higher than normal rainfall oc- 
curred in May and June (1978) but was spotty in distribution; general rains were not 
experienced at this time. Rainfall in July was generally below normal, while August was 
wetter than normal. 

The scattered nature of the 1978 rainfall (in both time and space) resulted in a mosaic 
of areas with flush growth of the larval food plant, C. pallida. Isolated centers of butterfly 
concentrations developed and generated the comparatively local migrations which were 
observed in July and September 1978. 

The lack of migrating snout butterflies in deep southern Texas was due to two factors. 
Lack of butterflies in the Llano Mesteno area was due to lack of the prime larval food- 
plant. Lack of butterflies south of the Llano Mesteno in the Lower Rio Grande Valley 
(area along the Rio Grande, including Brownsville) was due to lack of heavy rainfall. 
Continued drought .in this latter area precluded rapid growth of the prime larval food- 
plant, C. pallida. 

I thank George Toalson and J. Stephen Neck for observations on the movements of 
snout butterflies in 1978. T. B. Samsel III drafted Fig. 1. 


RAYMOND W. NECK, Texas Parks and Wildlife Department, 4200 Smith School Road, 
Austin, Texas 78744. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 323 


A BILATERAL GYNANDROMORPH OF ANTEPIONE THISOARIA 
(GEOMETRIDAE) 


An interesting bilateral gynandromorph of the geometrid moth, Antepione thisoaria 
(Guenée), was collected at mercury vapor light in August 1974 about 11 km southwest 
of Nashville, Tennessee. This specimen (Fig. 1) has a wingspan of 34 mm with male 
hindwing coloration on the right side and female to the left; the irregular brown patches 
towards the margins of each forewing suggest a mixture of male and female traits. The 
wing-coupling apparatus and genitalia are of male type on the right side and of female 
type to the left. 

I am grateful to A. Watson and Drs. D. S. Fletcher, R. W. Hodges and D. C. Ferguson 
for their comments concerning this specimen which is presently in the author’s possession. 


LANCE A. DURDEN, Department of Anatomy, Vanderbilt University School of Med- 
icine, Nashville, Tennessee 37282. 





Fic. 1. Bilateral gynandromorph of Antepione thisoaria (Guenée). 


Journal of the Lepidopterists’ Society 
38(4), 1984, 324-3827 


BOOK REVIEWS 


Systematische Untersuchungen am Pieris napi-bryoniae-Komplex (Lepidoptera: Pieri- 
dae), by Ulf Eitschberger. 1983. Herbipoliana 1(1), 504 pp.; 1(2), 601 pp. Published by 
the author and Hartmut Steiniger. Available from the author at Humboldtstrasse 13, 
D-8671 Marktleuthen, West Germany, and from entomological book dealers. (Due to 
fluctuations of the Mark against the Dollar, current prices are not available. Early in 
1984 the price was 360.- DM + 15.— DM international postage. ) 


The biosystematics of the Pieris napi group remains one of the great intractable 
problems in the Holarctic butterflies. This is despite the massive and valiant revisionary 
effort represented by this lavishly-produced monograph, which has truly been a labor of 
love for its author. 

Systematische Untersuchungen (“Systematic Investigations’), hereafter referred to as 
S.U., brings together in one place more morphological and distributional data on the 
napi group than have ever been assembled before. Eitschberger did an incredible amount 
of finely detailed morphological work, which is reflected in extraordinary series of pho- 
tographs (optical and SEM) and drawings of characters of both adults and immatures. 
The entire second volume is made up of illustrations, among them 218 color plates 
averaging over 30 specimens/plate. The photographs are meticulously produced and the 
colors by and large very true. Within each taxon a range of variability is usually repre- 
sented, including seasonal forms and sexual differences; for napi and bryoniae numerous 
aberrations, rare genetic morphs, and sexual mosaics are also presented. The same spec- 
imens are usually shown in upper and lower surfaces on the same plates. For some 
reason the ventral surfaces are printed slightly smaller than the corresponding dorsals, 
which is confusing. There is no back-referencing system from the plates to the text, and 
the forward-referencing system is somewhat clumsy. 

The first volume of S.U. contains all the text, plus numerous distributional maps. 
Except for long quotes from the primary literature, which are reproduced from the 
originals by photo-offset and are thus in their original languages, the text is in German 
and will not be easy going for readers unskilled in that language. (The most important 
previous work on the group, the monograph on napi and bryoniae by Miiller and Kautz, 
is also in German and is even more strenuous reading. Moral: If you want to work on 
the napi group, learn German.) 

Volume 1 is divided into a fairly brief overview of previous taxonomic work in the 
group and of the morphological characters deemed to be of value in such work, and a 
very lengthy taxon-by-taxon treatment which does include “biological,” live-bug infor- 
mation when available. Twenty-five species are recognized, with a total of 48 subspecies 
in addition to the nominate ones. The species are grouped into four sets: a Eurasiatic 
complex of 11 species, including true napi and bryoniae; a North American group of six 
(to be discussed below); and two Asiatic groups of four species each. There is a tabular 
summary of character states for the taxa of the first group (pp. 46-51). 

Even a casual inspection of volume 1 reveals a number of potential problems. (1) 
Geographic coverage is extremely uneven. This is presumably no fault of the author, 
who in fact has been remarkably successful in assembling material from odd places. But 
as one might expect, distributions are mapped in almost infinitesimal detail in western 
Europe (diminishing rapidly to the east!), moderate (and to this reviewer, rather unsat- 
isfactory) detail in North America, and poorly indeed in Asia—where, except for Japan, 
most taxa are represented by a handful of widely separated, random-looking dots on the 
map. The inevitable result is that taxonomy is much coarser in some areas than in others. 
(2) The author is not an ideologue, and does not attempt to force the taxa into the 
formalisms of cladistics or the quantitative definitional modes of phenetics. He is, how- 
ever, apparently not much of an evolutionist or biogeographer either, and he has an old- 
fashioned, implicitly typological and explicitly morphological species concept. His work 
thus most resembles the alpha-taxonomy done on poorly-known groups of bark beetles 
from Java, and is not at all like what one has come to hope for in the Holarctic butterflies 


VOLUME 38, NUMBER 4 320 


in these sophisticated times. (3) The naming of new taxa has been promiscuous and based 
on the sort of species concept just described. Many of the new taxa are unlikely to sit 
well with regional specialists, and many are apt to be ignored or to be treated as junior 
synonyms of more familiar names, at least until more information about the biology of 
the animals is available. All these points are relevant to the handling of the Nearctic 
fauna. 

Eitschberger recognizes 18 taxa in the Nearctic, of which nine, or 50%, are new. They 
are (* = Eitschberger name): Pieris venosa venosa; P. oleracea oleracea; P. o. ekisi*; P. 
marginalis marginalis, P. m. reicheli*; P. m. pallidissima; P. m. mcdunnoughi; P. m. 
mogollon; P. m. hulda; P. m. meckyae*; P. m. guppyi*; P. m. tremblayi*; P. m. shapiroi*; 
P. m. browni*; P. acadica acadica; P. angelika angelika*; P. virginiensis virginiensis; P. 
v. hyatti*. He is not certain that all the marginalis subspecies are conspecific. There are 
also brief discussions of several additional marginalis populations he is unwilling to name 
for lack of good series. Most of the new taxa occur in northwestern North America, from 
Alaska to British Columbia. (P. angelika, named for Eitschberger’s wife, was actually 
described in 1981 in a paper in the German journal Atalanta, which Eitschberger edits. 
It is generally unheard of in North American lepidopterological circles. The other new 
taxa are named and described in S.U. itself. Angelika is described at the species level 
for reasons which are not terribly clear. It is mostly allopatric with the various marginalis- 
taxa, but there is a suggestion of sympatry in a few places. Aside from the co-occurrence 
of oleracea and virginiensis in a few localities in the northeastern U.S. and perhaps 
adjacent Canada, this would be the only instance in which it is alleged that members of 
the napi complex occur sympatrically in the Nearctic.) 

I have accused Eitschberger of being typological, and I should qualify this by saying 
that a summary of character-state distributions—raw data only—for selected wing char- 
acters is given for most taxa based on the series he examined; and, as noted already, the 
illustrations portray a good range of variation. Nonetheless, one is left unsatisfied as to 
the criteria used to recognize and rank taxa; basically, we are being asked to trust the 
author’s judgment. I have discussed this with Eitschberger with specific reference to the 
northwestern Nearctic taxa, and it is quite plain to me that his weighting criteria are 
perfectly clear to him. But they are not to me. In fact, I do not consider my own 
patronymic, shapiroi—which I have never seen alive; oddly, I have apparently worked 
on the population Eitschberger named angelika—to be well-defined and find it a good 
candidate for sinking. (I won't miss it.) 

North Americans tend to bristle at the idea of Europeans working on their fauna from 
a distance; after all, we are no longer colonials. Such jingoistic reactions should play no 
role in how we evaluate Eitschberger’s treatment of the Nearctic napi. Most Nearctic 
workers who know our taxa by experience will, however, be properly suspicious of his 
weighting and grouping. Northern California workers, for example, know that a very 
complex situation exists in that region in which venosa, marginalis, and pallidissima are 
all involved; there is no hint of that here. Entities which are considered allospecific by 
Eitschberger may or may not be interbreeding in such zones. Such information must 
ultimately override inferences from morphology. Eitschberger’s logical structure would 
fall apart if interbreeding cuts across his morphological criteria for species status. But 
does it? 

The pitfalls, not only of Eitschberger’s methods but of their application to this partic- 
ular group, are shown by his contribution to the seemingly endless European napi- 
bryoniae problem. These two taxa are to European butterfly work what Colias philodice 
and eurytheme are in North America. Are they one species, or two, or something some- 
how inbetween? Given that they appear to interbreed in some places but not others, Z. 
Lorkovié proposed that they be treated as “semispecies,” species in statu nascendi. But 
“semispecies’ is not a taxonomic ranking, and one must decide what to call them. After 
much soliloquizing, Eitschberger opts to treat them as species and indeed to give bryoniae 
fifteen (!) subspecies of its own—extending the sense of the name to a large number of 
poorly-known, hitherto obscure, and very interesting Asiatic populations. 

Meanwhile, at Bern, Switzerland, Hansjiirg Geiger (1978, Entomol. Zeitschrift 88:229- 
235; 1981, J. Res. Lepid. 19:181-195; 1985, Experientia, 41:24—29) has shown that elec- 


326 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


trophoretically European napi and bryoniae are virtually identical (within-taxon variance 
sometimes exceeds between-taxon variance). This of course does not prove conspecificity 
(see below). It is entirely consistent with the hypothesis that the napi-bryoniae distinction 
is very recent (Holocene) as compared to many other taxic distinctions in Pieris, and it 
seems inconsistent with the schema developed by Ejitschberger. Speaking from the gut, 
I am willing to bet that electrophoretic data will show Eitschberger’s bryoniae-concept 
to be grossly polyphyletic. Again, time will tell. 

Why is all this so unsatisfactory? Part of the problem is that Eitschberger’s roots are 
in the German morphological tradition and not in Darwinism, so that we are not all 
speaking the same theoretical language. But that is only part of the problem; the other 
part derives from the animals themselves. The taxonomic characters in the napi-bryoniae 
group (if it is a monophyletic group) are mostly wing-color and -pattern things—things 
we know are commonly determined by a handful of loci. The correspondence between 
these characters and reproductive isolation is not good, as Lorkovié has shown us with 
his sibling species balcana. Most revisionary work in Lepidoptera involves morphological 
characters, especially genetalic ones. But in this group these characters are close to 
worthless. The differences are so slight—as the hundreds of drawings in vol. 2 show us— 
that Eitschberger is forced to seize on trivia and to weight them heavily in order to be 
able to define taxa at all. The results are, not unsurprisingly, not very satisfactory when 
we compare the napi complex to other groups in which nature has been kinder to the 
taxonomist. 

And that is not all; such differences as exist are often overshadowed by phenotypic 
plasticity in the napi group. Most of the taxa are polyphenic; seasonal phenotypic dif- 
ferences within taxa are normally greater than differences in the corresponding seasonal 
brood between taxa; and pupal morphology varies depending on whether or not the 
individual is in diapause. All of this spells a mess of overwhelming proportions. 

I maintain that in many cases, only reproductive-compatibility and/or electrophoretic 
data will permit proper species assignments in this group. Yet we can now see that these 
two types of data may conflict with each other! Geiger finds that, electrophoretically, 
balcana is no more different from napi than bryoniae is—but balcana and napi are 
highly intersterile (Lorkovié) while bryoniae and napi are more or less highly interfertile, 
with some exceptions. There is a hint in all of this of an infective or transposable genetic 
element responsible for sterility, something akin to the “hybrid dysgenesis” factors in 
Drosophila. If that should be the case, it kicks all conventional species concepts into a 
cocked hat. 

Whatever the basis of sterility in hybrid crosses, it is certain that character reversals, 
parallelisms, and the like are lurking everywhere in the napi group. The European group 
of taxa clustering electrophoretically with napi, and probably the Nearctic ones grouped 
in marginalis and angelika by Eitschberger, seem to represent rapidly-evolving com- 
plexes consequent on Pleistocene and post-Pleistocene events. In northwestern North 
America there is reason to think one is dealing with two, perhaps three, invasions across 
Beringia—the oldest perhaps Tertiary, the youngest very recent. Preliminary electro- 
phoretic data from Geiger, not available to Eitschberger, roughly support the latter’s 
gross clustering of Nearctic taxa. 

No grand synthesis is possible without genetics or without historical-biogeographical 
analysis or without cladistic reasoning (if not cladistic formalisms). If and when all this 
is done—and we intend to try it for the northwestern Nearctic—I suspect Eitschberger’s 
judgments will be shown better than might have been forecast. But, I do not intend to 
start using his nomenclature until then. 

Lionel Higgins has given this book a favorable review in the British journal Entomol- 
ogist’s Gazette (1984, 35:174-175). I cannot be so sanguine. (Higgins, by the way, seems 
to have gotten lost in this Teutonic tome; he asks what the structures shown in vol. 2, 
pp. 317-321, might be. They are the distal ends of the tongue-cases and antennae of 
pupae, as should be self-evident to anyone who has seen pierine pupae but at any rate 
is explained in vol. 1, p. 15.) Eitschberger, who is by profession a pharmacist, has gone 
to incredible trouble and almost unimaginable personal expense to provide us with this 
huge work. As I noted at the beginning, it is the biggest compilation of data on these 


VOLUME 38, NUMBER 4 327 


animals ever assembled, and therein lies its primary value. The data cry out for other 
sorts of interpretation than Eitschberger has given them. Anyone interested in this most 
exasperating of groups, and who reads German, must have access to this book. If you 
are not willing to buy it, have your institutional library do so. Otherwise, it will become 
another Miiller and Kautz. Possibly the only academic library copy of Miiller and Kautz 
in the United States is at Yale, which prohibits photocopying of interlibrary loan mate- 
rials. The only way to get hold of the book is to go to New Haven or to buy one through 
an antiquarian. Will S.U. disappear in similar fashion? 

Eitschberger told me over a stein of beer that he hopes other people will take up and 
expand his work. That is good, for it must—and will—be done. Systematische Unter- 
suchungen ... could be read superficially (and apparently was, by Higgins) as the de- 
finitive resolution of the napi problem. It isn’t. It is a beginning. 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 327-328 


DEAR LORD ROTHSCHILD (BIRDS, BUTTERFLIES & HISTORY). Miriam Rothschild. 1983. 
Hutchinson Publishing Group, 17-21 Conway St., London, W1P5HL. Format 7” x 934” 
398 pp., including index & appendices. 90 pp. of B/W photographs. 12 pp. of color 
plates. Cloth bound. 14.95 (British pounds.) 


One usually thinks of Lord Rothschild in connection with Karl Jordan or Ernst Hartert, 
both of whom were among his co-authors. In contrast, few people know the history of 
the Tring Museum, nor the other aspects of his life which took place beyond the bound- 
aries of Tring. This book, written by his niece, is a revelation. It is a story of one life, 
liberally embellished with ancestors and heavily endowed with wealth. It is the story of 
the workings of Parliament, the education of the young, of action on the high seas, and 
of wild creatures, both alive and dead, which previously had never been known to the 
world. 

“Tt is not easy to be born. The average man is squeezed out into the world with blood 
to lubricate his passage and wild shrieks of anguish to speed him on his way.” 

So begins the biography of Lionel Walter Rothschild, 2nd Baron of Tring. 

At the age of seven, at tea time in the nursery, Walter suddenly stood and made the 
following announcement: “Mama, Papa, I am going to make a museum, and Mr. Minal 
is going to help me look after it.’ This prophesy came true. 

Walter was the classic example of a child who shows little scholastic promise, but at 
some point becomes fired with enthusiasm in one particular field of endeavor to the 
extent that he becomes expert in that field to the exclusion of all else. A psychologist 
might have altered this, if such a person had existed in England at that time. He was 
born to a mother who was strict and sensorious on one hand—overprotective and indul- 
gent on the other. His father was never able to understand either his love for animals or 
his failure in finances. From the beginning he had a speech defect which resulted in 
crippling shyness. He was tutored at home and rarely played with boys of his own age. 
All of this, added to the astronomical wealth of his family, contributed to his enigmatic 
personality. 

He began his collections at the age of seven with one butterfly. By the time he was 19 
he had collected 5000 birds (2000 of which he had already mounted) and 38,000 Lepi- 
doptera. Two years later, his family built him a museum as a 21st birthday present. 

This book mirrors the life-long curiosity of one man to discover and collect all the 


328 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


exotic forms of life, many of which are now extinct, or nearly so, from immense animals 
such as the Galapagos tortoises and elephant seals to the smallest of insects, the “fleas 
and lice from bats, birds and mammals.”’ Included in his final inventory were 13 Gorillas, 
62 Birds of Paradise, 520 Hummingbirds, 144 Giant Tortoises, 300,000 bird skins, 200,000 
birds eggs and 2,250,000 Lepidoptera. Obviously such carnage would not be tolerated 
today, but perhaps it was a necessary prelude to the conservation movement, of which 
Charles Rothschild is considered to be the founder. In any case, the variety of perfectly 
mounted animals and birds were, at that time, a revelation and a major contribution to 
the understanding of the animal kingdom. 

In 1889 when he was 21, Walter entered the firm of N. M. Rothschild & Sons, at the 
insistance of his father. He resigned in 1908, having spent most of his tenure not in 
banking but in editing his museum’s publication, “Novitates Zoologicae,” writing mono- 
graphs and, with his salary, hiring collectors, planning and financing expeditions to 
remote places and corresponding with his collectors and crews. 

During this same period he became involved with two scheming women at the same 
time, and still worse, his antics with them were such that an unidentified (and financially 
deprived) peeress enjoyed a life-long income by periodically blackmailing him. The First 
World War depleted his fortune still further. 

His last decisive act was to sell his cherished collection of birds to the American 
Museum of Natural History in New York. He never told his family what he had done, 
and they only found out through an article in the Times five months later. After this 
deeply traumatic sacrifice, his ardour diminished. He died five years later. 

The author’s obvious esteem for her uncle is felt throughout the book. She narrates 
the adventures and the misadventures of this enigmatic man without censure, without 
apology, but with sensitivity and candor. She seems to lift him out of her feelings and 
set him in a place in the sun where all of his many facets can glisten and come and go. 

The illustrations are outstanding. They are nearly a biography in themselves. Walter 
is seen at all ages, from adorable to cute to beautiful, to handsome, to distinguished. His 
mother is pictured in her coronation robes, worn at the coronation of Edward VII. There 
are many pictures of Tring Palace, Tring Park, ancestors and relations, one of King 
George V at Tring, and one of Queen Victoria—autographed. 

The Victorian era has never seemed more wondrous than in the two generations of 
Rothschilds which dominate this bbok—Nathan Mayer, First Baron Rothschild of Tring 
and his wife, Emma Louise von Rothschild and their two sons, Lionel, Second Baron 
Rothschild of Tring and Nathaniel Charles. Hon. Miriam Rothschild, the author, is a 
daughter of Charles. It is owing in large part to her sympathy that this extraordinary 
family is made so lovable. In the end one has not read the story of one man only but of 
a devoted family whose fabulous wealth was shared by the world in ways not always 
clear, nor even comprehensible, but certainly in this book, memorable. 


Jo BREWER, 257 Common Street, Dedham, Massachusetts 02026. 


Journal of the Lepidopterists’ Society 
38(4), 1984, 329-332 


INDEX TO VOLUME 38 


(New names in boldface) 


Acanthopteroctetes aurulenta, 47 
Acer negundo, 318 
Acroceras zizanioides, 104 
Actias luna, 116 

A. maenas, 114 

A. selene, 118 

A. sinensis heterogyna, 117 

A. truncatipennis, 118 
Adams, E. R., 318 
Adamski, D., 317 
Adinandra dumosa, 117 
Agapema galbina anona, 137 

A. homogena, 134 
Agraulis vanillae, 28 
Agriphila ruricolella, 150 

A. vulgivagella, 150 
Amphipyra tragopoinis, 317 
Andersen, W. A., 63 
Anisota consularis, 143 

A. oslari, 51 

A. senatoria, 51 
Antepione thisoaria, 323 
Anthocharis lanceolata, 251 
Apateticus cynicus, 61 

A. lineolatus, 61 
Apterona helix, 317 
Arbogast, R. T., 202 
Areca catechu, 78 
Argema mimosae, 118 

A. mittrei, 118 
Argyria nivalis, 151 
Arnolds J.-R:, 257 
Asclepias spp., 209 
Asterocampa celtis, 253 

A. clyton, 186, 253 

A. clyton flora, 186 


Asterocampa spp. (parasites & predators): 


6 

Atlides halesus, 179 
Atrichum undulatum, 51 
Autographa flagellum, 92 

A. rubida, 95 
Automeris, 281 
Averrhoa bilimbi, 117 
Basacallis, 275 

B. tarachodes, 268 
Battus philenor, 142 
Becker, V. O., 18 
Blanchard, A., 245 
Book Reviews: 68, 147, 254, 324, 327 
Brachymeria sp., 61 
Brassica campestris, 243 

B. geniculata, 244 


B. Kaber, 243 
B. nigra, 243 
Brewer, J., 327 
Brow! V2Ac. jr. 96 
Brown, J. W., 138 
Brown, L. N., 65 
Byrd, R. V., 202 
Caerois sp., 103 
Caligo atreus uranus, 103 
C. memmon, 103 
Callosamia angulifera, 261 
C. promethea, 265 
C. securifera, 265 
Cameron, E. A., 57 
Carex spissa, 138 
Cargida pyrrha, 88 
Carrie W261) 
Caryota rumpha, 78, 81 
Cashatt, E. D., 268 
Castilleja linariaefolia, 9 
Celtis laevigata, 186 
Chauvin, G., 202 
Chrysoteuchia topiaria, 150 
Cocos nucifera, 81 
Coleophora laricella, 235 
Colias eurytheme, 67 
Coloradia pandora, 65, 281 
C. p. lindseyi, 284 
C. velda, 284 
Condalia lycioides, 88 
Conecephalum conicum, 41 
C. conium, 192 
Cordyline terminalis, 78 
Cotesia spp., 60 
Crambus ainslieellus, 150 
. alboclavellus, 150 
. coloradellus, 150 
. laqueatellus, 150 
. leachellus, 150 
. pascuellus floridus, 150 
. perlellus innotatellus, 150 
. praefectellus, 150 
Cycas circinalis, 70 
Cynodon dactylon, 138 
Danaus gilippus, 148 
D. plexippus, 209 
Davis DD. Re 47 
Deschampsia caespitosa, 138 
Durden, L. A., 323 
Ecpantheria deflorata, 192 
Ehrlich, P. R., 1 
Eichlin, T. D., 18 
Elachertus sp., 60 


OO AOE) GO 


330 


Elymnias agondas, 83 
Emarginea percara, 184 
Endrosis sarcitrella, 203 
Eoreuma crawfordi, 151 
Epiblema luctuosana, 245 
E. luctuosissima, 245 
Epimartyria pardella, 40 
Eucalyptus gunnii, 117 
Euchloe ausonides, 242 
Euchromius californicalis, 151 
Euphorocera floridensis, 60 
Euphydryas gillettii, 1 
Evans, D. L., 194 
Fauske, G., 149 
Ficus calopilina, 22 
F. semivestita, 22 
Flowers, R. W., 139 
Foeniculum vulgare, 195 
Formica integra, 124 
Friedlander, T. P., 60, 139 
Galleria mellonella, 207 
Gillaspy, J. E., 142 
Glaucopsyche lygdamus, 124 
Godfrey, G. L., 88 
Godyris zavelata caesiopicta, 62 
Graellsia isabellae, 116 
Gustin, R. D., 149 
Hamadryas amphichloe diasia, 173 
H. a. ferox, 174 
H. amphinome mexicana, 172 
H. atlantis lelaps, 173 
H. februa ferentina, 172 
H. feronia farinulenta, 172 
H. fornax fornacalia, 174 
H. guatemalena marmarice, 173 
H. iphthime joannae, 173 
Helianthus sp., 92 
Heliconius charitonia, 141 
Heliothis phyloxiphaga, 310 
Hemileuca burnsi, 284 
. chinatiensis, 282 
. diana, 284 
. eglanterina annulata, 283 
e. eglanterina, 283 
e. shastaensis, 283 
electra clio, 284 
e. electra, 284 
griffini, 282 
hera hera, 283 
h. marcata, 283 
juno, 284 
lucina, 51 
neumoegeni, 284 
. nevadensis, 51, 284 
. nuttalli uniformis, 283 
. tricolor, 284 
Heppner, J. B., 68 


be BB SB ge 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Herrania albiflora, 249 
Hochberg, M. E., 176 
Hoffman, L. R., 192 
Hofmannophila pseudospretella, 202 
Holdren, C. E., 1 
Holland, R., butterflies of two New Mexico 
mountains, 220 
Hookeria lucens, 41 
Humiphila paleolivacea, 276 
Hyalophora cecropia, 261 
Hyantis hodeva, 82 
Hypolimnas deois, 83 
Hypothyris euclea leucania, 62 
Immature Stages (descriptive): 
Ova, 2, 115, 18, 19) 7OM7OROARII2 04: 
240, 243, 287, 288, 312 
Larva, 2, 15; 18) 19) 41) o2eoAaON iG: 
79, 89, 92, 104, 121, 134, 240, 248, 
287, 288, 306, 312 
Pupa, 4, 16, 18, 19, 54, 70, 76, 80, 106, 
WAP BLP? 
Itoplectis conquisitor, 61 
Jenkins, D. W., 171 
Juniperus virginia, 144 
Khalaf, K. T., 64 
Kitching, R. L., 209 
Klassen, P., Manitoba butterfly checklist, 
32 
Lamarkia aurea, 138 
Lambremont, E. N., 252 
Lara, J. R., 142 
Larix decidua, 235 
L. occidentalis, 235 
Leffler, S. R., 235 
Lespesia aletiae, 60 
Matrisisp 92 
Libytheana bachmanii, 139, 319 
L. carinenta, 139 
Liquidambar formosana, 118 
L. styraciflua, 117 
Liriodendron tulipifera, 262 
Moereh Ga Reon 
Lonicera involucrata, 4 
Luthrodes cleotas, 72 
Lymantria dispar, 57 
Macaranga aleuritoides, 16 
M. involucrata, 18 
M. quadriglandulosa, 16 
Madia glomerata, 310 
Malaisia scandens, 20 
Manduca morelia, 96 
M. pellenia, 96 
M. wellingi, 96 
McDaniel, B., 149 
Mechanitis spp., 61 
Megalopyge opercularis, 64 
Melinaea spp., 61 


VOLUME 38, NUMBER 4 


Melittia calabaza, 13 
M. cucurbitae, 18 
M. pauper, 14 
Metadontia amoena, 183 
Meteorus spp., 60 
Microcharops tibialis, 60 
Microcrambus elegans, 150 
Mitoura gryneus, 144 
Morpho granadensis polybaptus, 103 
M. peleides limpida, 103 
Morphopsis albertisi, 69 
Mulford, B. L., 310 
Musa sp., 78 
Mycalesis drusillodes, 83 
Myscelia ethusa, 103 
Nassig, W. A., 114 
Neck, R. W., 319 
Neil, K., 92 
Nessaea aglaura, 103 
Nielsen, M. C., 124 
Obituaries: 257, 259 
Occidentalia comptulatalis, 151 
Opler, P. A., 147 
Pandanus sp., 81 
Papilio aegeus, 83 
. antinous, 165 
. eurymedon, 165 
. glaucus, 165 
. machaon syriacus, 194 
. turnus, 165 
. victorinus, 237 
Parachma borregalis, 268 
P. ochracealis, 268 


sof loys} Tavs) Tonsl Ise} Ive) 


Parasites of Lepidoptera: 9, 60, 119, 183 


Paratrytone melane, 138 
Parsons, M., 15, 69 
Pediasia dorsipunctella, 151 

P. luteolella, 151 

P. mutabilis, 151 

P. trisecta, 151] 
Pedicularis bracteosa, 9 
Peigler, R. S., 51, 114 
Pellia sp., 41 
Phaius tancarvilleae, 78 
Philiris agatha, 18 

P. helena, 15 

P. intensa, 18 

P. moira, 15 

P. ziska, 19 
Phoradendron tomentosum, 179 
Phryganidia californica, 176 
Pieris rapae, 66 
Pinus contorta, 65 
Piper, G. L., 310 
Pipturus argenteus, 19 
Plantago lanceolata, 192 

P. rugellii, 192 


331 


Platytes vobisne, 151 
Podistes exclamans, 61 
Podisus maculiventris, 61 
Predators of Lepidoptera: 61 
Prepona spp., 103 
Prunus virginiana, 265 
Ptermalus vanessae, 9 
Quercus agrifolia, 125 
Q. alba, 176 
Q. coccinea, 54, 125 
Q. velutina, 125 
Rhamnus californica ursina, 136 
Rhus cipallina, 117 
R. glabra, 117 
R. radicans, 117 
R. typhina, 117 
Rutowski, R. L., 23 
Salix gracilis, 52 
Samia cynthia, 266 
Saturnia, 281 
Satyrium edwardsii, 124 
S. kingi, 68 
Schaefer, J., 23 
Schima wallichii, 117 
Schizura rustica, 245 
Seigler, D. W., 192 
Sevastopulo, D. G., Gesneriaceae & Big- 
noniaceae as food-plants, 235 
Sevastopulo, D. G., food-plants of Pieridae, 
249 
Shapiro, A. M., 147, 242, 251, 324 
Shuey, J. A., 144 
Similia camelus, 126 
Similipepsis aurea, 85 
S. ekisi, 86 
S. lasiocera, 85 
S. typica, 85 
S. violaceus, 85 
Sinea sanguisuga, 61 
S. spinipes, 61 
Smilax sp., 81 
Smith, M. J., 134 
Smith, N. J., 40 
Sorensen, J. T., 254 
Spencer, K. C., 192 
Stamp, N. E., 186 
Stenotaphrum secundatum, 138 
Streptanthus howellii, 251 
Strong, R. G., 202 
Synanthedon acerrubri, 318 
Taenaris artemis, 81 
T. bioculatus, 83 
T. butleri, 72 
T. catops, 70, 76 
T. dimona, 81 
T. gorgo, 81 
T. horsfieldii, 70 


332 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


T. mailua, 82 Townsend, R. F., 259 

T. myops, 79 Turpinia sphaerocarpa, 117 

T. onolaus, 70 Tuskes, P. M., 40, 134, 281 

T. phorcas, 70, 81 Tuttle, J., 143 
Tapenochilus sp., 79 Tyler, H. A., 257 
Taygetis andromeda, 102 Upton, M. S., 165 

T. ypthima, 112 Valeriana occidentalis, 9 
Telenomus spp., 60 Vespula sp., 61 
Tetrastichus spp., 60 Volney, W. J. A., 176 
Thaumatopsis fernaldellus, 151 Wang, P. Y., 85 

T. pectinifer, 151 Webster, R. P., 124 

T. pexellus, 151 Westcott, R. L., 259 

T. repandus, 151 Whittaker, P. L., 179 
Theclinesthes onycha, 72 Williams, B. D., 51 
Theobroma cacao, 249 Williams, E. H., 1 
Thopeutis forbesellus, 151 Wourms, M. K., 67 
Tinea pellionella, 208 Xanthopimpla spp., 119 
Tineola bisselliella, 208 Young, A. M., 61, 65, 102, 141, 237, 245 
Tisiphone helena, 69 Zalucki, M. P., 209 


Date of Issue (Vol. 38, No. 4): 10 July 1985 


EDITORIAL STAFF OF THE JOURNAL 
THoMaS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MacDaA R. Papp, Editorial Assistant 
DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
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plant or animal in the text should include the full scientific name, with authors of 
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Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


HYBRIDIZATION BETWEEN CALLOSAMIA AND HYALOPHORA 


(SATURNIIDAE). Thomas W. Carr... ee 261 
REVISION OF THE GENUS PARACHMA WALKER (PYRALIDAE: 
CHRYSAUGINAE) OF NORTH AMERICA NORTH OF MEXICO 
WITH DESCRIPTION OF A NEW GENUS. Everett D. Cashatt 268 
THE BIOLOGY AND DISTRIBUTION OF CALIFORNIA HEMILEUCIN- 
AE (SATURNIIDAE).. Paul M. Tuskes .__ a 281 
OBSERVATIONS ON THE BIONOMICS OF HELIOTHIS PHYLOXIPHAGA 
(NOCTUIDAE) ON CLUSTER TARWEED IN SOUTHEASTERN 
WASHINGTON. G. L. Piper & B. L. Mulford 33s 310 
GENERAL NOTES 
An unusual ovipositional site for Amphipyra tragopoginis (L.) (Noctui- 
dae). D> Adamskiu. 25 317 
Acer negundo (boxelder) as a food plant for Synanthedon acerrubri (Sesi- 
idae). . Eleaner R: Adams, 200 Eee 318 
On the origin of snout butterflies (Libytheana bachmanii larvata, Libythei- 
dae) in a 1978 migration in southern Texas. Raymond W. Neck .......... 319 
A bilateral gynandromorph of Antepione thisoaria (Geometridae). Lance 
Ay Dende yok ea OS 323 
BOOK REVIEWS). oputinn Wa Oe 0 Ol 324, 327 
INDEX TO; VOLUME 36g i008 Fl ee 329 


Volume 39 1985 Number 1 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





16 October 1985 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Don R. Davis, President LEE D. MILLER, 

Vi1TOR O. BECKER, Vice President Immediate Past President 
JAVIER DE LA MAZA E., Vice President JULIAN P. DONAHUE, Secretary 
JOHN C. DOWNEY, Vice President Eric H. METZLER, Treasurer 


Members at large: 


F. S. CHEW J. M. BURNS B. A. DRUMMOND 
G. J. HARJES F. W. PRESTON J. LANE 
E. H. METZLER N. E. STAMP R. K. ROBBINS 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
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Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$25.00 (institutional subscriptions) and $18.00 (active member rate) by the Lepidopter- 
ists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Boule- 
vard, Los Angeles, CA 90007. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, 
1900 John St., Manhattan Beach, CA 90266. 


Cover illustration: Micropylar end view (130) of the egg of Sericosema sp. (probably 
juturnaria) (Geometridae). The scanning electronmicrograph was taken by Thomas D. 
Eichlin, Sacramento, of eggs furnished by Ron Robertson, Santa Rosa, California. 


JOURNAL OF 


Tue LeEPIDOPTERISTS’ SOCIETY 














Volume 39 1985 Number 1 





Journal of the Lepidopterists’ Society 
89(1), 1985, 1-8 


NEW U.S. RECORDS AND OTHER INTERESTING 
MOTHS FROM TEXAS 


ANDRE BLANCHARD 
3023 Underwood, Houston, Texas 77025 


AND 


EDWARD C. KNUDSON 
808 Woodstock, Bellaire, Texas 77401 


ABSTRACT. Twenty-eight moths, most of which are recorded from the U.S.A. or 
Texas for the first time, are illustrated. The text includes a brief description and distri- 
butional records known to the authors. 


This paper reports 28 species of moths collected in Texas by the 
authors. In most cases, these have not been illustrated previously or 
have been mentioned only in publications that are now long out of 
print and difficult to obtain. The species were selected because in some 
cases they represent new U.S. records and are not included in the new 
Check List of the Lepidoptera of America North of Mexico (Hodges 
et al., 1983); in other cases, the species were included in the new check 
list based on the records given here, the specimens having been pre- 
viously examined by the authors of the check list. Certain species were 
included because they represent important new range extensions. Fi- 
nally, some represent species that have not been adequately illustrated 
previously. The authors hope that this information on striking and 
easily recognized species will be useful and that perhaps it will en- 
courage others to develop an interest in this field of lepidopterology. 


Sphingidae 


Xylophanes libya Druce (Fig. 1). Hidalgo Co., Bentsen Rio Grande Valley St. Pk., 11- 
X-75, 1 male, E. Knudson coll. New U.S. record. Det. R. Hodges and V. Brou. This 
species is common in Mexico and has been collected there by the junior author within 
200 miles of the border. The forewings are several shades of olive brown; hindwings are 
banded with black and dull orange yellow. 


2 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Noctuidae 


Acronicta funeralis (G. & R.) (Fig. 2). Washington Co., Brenham, 1-IV-79, 2 males (1 
donated to USNM), 1 female, 1 last instar larva on Hickory (Carya sp.), producing a 
male, May 1980; Anderson Co., Engeling Wildlife Management Area, 15-III-83, 2 males, 
all E. Knudson coll. Det. R. Poole, These are apparently the first Texas records of this 
species. Forewings light gray marked with black; hindwings white. 

Euxoa pimensis B. & McD. (Fig. 3). El Paso Co., Tom Mays Park, 24-V-81, 3 males 
(1 donated to CNC), 3 females; Culberson Co., Noel Creek, 26-V-81, 1 male, all E. 
Knudson coll. Det. Lafontaine. Apparently a new Texas record. Forewing light gray with 
variable black shading between orbicular and reniform, black basal dash; hindwings pale 
fuscous. 

Eriopyga iole Schaus (Fig. 4). Brewster Co., Big Bend Nat'l. Park, Green Gulch, 28- 
V-81, 1 male, E. Knudson coll. New U.S. record. Det. R. Poole. Head, thorax and 
forewings reddish brown; ordinary lines obscure, dark brown; reniform outlined with 
white; hindwings pale fuscous, fringe orange-brown. 

Oncocnemis rosea Smith (Fig. 5). El Paso Co., Tom Mays Park, 30-III-83, 5 males, 6 
females (1 pair donated to USNM), E. Knudson coll. Det. R. Poole. Apparently, first 
Texas record for this species, which, with the preceding Euxoa pimensis, is from the 
infrequently collected Franklin Mountains near E] Paso. Forewings pinkish, shading to 
ochreous near base; strong black antemedial line; hindwings fuscous, lighter toward base. 

Oncocnemis terminalis Smith (Fig. 6). Hemphill Co., Lake Marvin, 9-X-82, 1 male, 
E. Knudson coll. Det. R. Poole. Probable new Texas record. Forewings dark brown with 
blackish markings; hindwings white, with broad black terminal band, fringe white. 

Miracavira brillians Barnes (Fig. 7). Brewster Co., Big Bend Nat'l. Park, Green Gulch, 
27-VI-65, 2 males; Chisos Basin, 25-VIII-65, 5 males, A. & M. E. Blanchard coll.; Chisos 
Basin, 10-VIII-83, 2 males, E. Knudson coll. Det. J. G. Franclemont. Forewing white and 
different shades of foliage-green, with darker pattern of deep brownish black; reddish 
brown tornal spot; hindwings white. 

Letis xylia (Guenée) (Fig. 8). Calhoun Co., Port Alto, 16-X-60, 1 male; Kleberg Co., 
Padre Island Nat'l. Seashore, 19-VII-76, 1 female, A. & M. E. Blanchard coll. New U.S. 
record. Det. E. Todd. The illustrated female is definitely paler than the male. Color is 
brown varying in saturation from very light to very dark. 

Cropia ruthaea Dyar (Fig. 9). Brewster Co., Big Bend Nat'l. Park, Green Gulch, 28- 
V-81, 1 male, E. Knudson coll. New U.S. record. Det. R. Poole. Forewings dark brown 
with black antemedial and postmedial lines; subterminal line, orbicular, and reniform 
contrasting light brown; hindwings light brown, paler toward base. Similar to Cropia 
connecta Smith, which also occurs in western Texas. Connecta is larger; has a contrast- 
ingly dark shaded inner margin of forewing; orbicular and reniform concolorous with 
ground; subterminal line indistinct; hindwing darker. 

Hemispragueia idella Barnes (Fig. 10). Presidio Co., Ruidosa Hot Spring, 8-VII-69, 1 
male, A. & M. E. Blanchard coll. Forewings ivory-white with black markings; hindwing 
bright yellow. 

Lesmone fufius (Schaus) (Fig. 11). Hidalgo Co., Santa Ana Refuge, 13-IX-80, 3 males; 
Bentsen Rio Grande Valley St. Pk., 27-V-82, 1 female. Det. R. Poole. Forewings grayish 
brown with blackish orbicular, reniform, and costal wedge near apex; hindwings con- 
colorous. Sexes similar. 

Lesmone formularis (Geyer) (Fig. 12). Hidalgo Co., Santa Ana Refuge, 28-XI-75, 1 
female, A. & M. E. Blanchard coll.; Kleberg Co., Kingsville, 1-XI-75, 1 female, Shu Wong 
coll. (submitted by J. Gillaspy). Wings grayish brown varying in saturation from medium 
to dark; straight line from inner angle to apex in hindwing is orange, bordered internally 
by blackish. Sexually dimorphic. Males collected by junior author in Mexico have a strong 


— 


Fics. 1-7. 1, Xylophanes libya Druce, Hidalgo Co., Bentsen Rio Grande Valley St. 
Pk., 11-X-75, wing expanse 71 mm. 2, Acronicta funeralis (G. & R.), Washington Co., 


VOLUME 39, NUMBER 1 3 





Brenham, 1-IV-79, wing expanse 34.1 mm. 3, Euxoa pimensis B. & McD., El Paso Co., 
Tom Mays Park, 24-V-81, wing expanse 41.2 mm. 4, Eriopyga iole Schaus, Brewster 
Co., Big Bend Nat’l. Park, Green Gulch, 28-V-81, wing expanse 28.3 mm. 5, Oncocnemis 
rosea Smith, El Paso Co., Tom Mays Park, 30-III-83, wing expanse 28.8 mm. 6, Onco- 
cnemis terminalis Smith, Hemphill Co., Lake Marvin, 9-X-82, wing expanse 35 mm. 7, 
Miracavira brillians Barnes, Brewster Co., Big Bend Nat’. Park, Chisos Basin, 27-VI-65, 
wing expanse 35.8 mm. (Figures not at same scale. Full wing expanse given for each 
specimen. ) 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


blackish diagonal line from near base to apex in forewings; the diagonal line of the 
hindwings is blackish, without any orange shading. 

Eulepidotis addens (Walker) (Fig. 18). Hidalgo Co., Bentsen Rio Grande Valley St. 
Pk., 11-X-80, 1 female, E. Knudson coll.; Cameron Co., Brownsville, 9-XI-69, 1 male, A. 
& M. E. Blanchard coll. New U.S. record. Det. J. G. Franclemont and R. Poole. Fore- 
wings with dull yellow ground over basal %, interrupted by dark violet-brown semime- 
tallic bars; apical % dull violet-brown; hindwings brownish with blue “eyespot” near 
tornus. Male less colorful, with the yellow ground of forewing replaced by medium violet- 
brown. 

Goniocarsia electrica Schaus (Fig. 14). Brewster Co., Big Bend Nat'l. Park, Green 
Gulch, 7-X-66, 1 female, A. & M. E. Blanchard coll. Det. E. Todd with the remark 
“extremely variable’ (pers. comm.). Medium brown with violaceous tinge; postmedial 
line whitish. 


Notodontidae 


Nystalea collaris (Schaus) (Fig. 15). Cameron Co., Brownsville, 8-XI-69, 1 female; 17- 
XI-69, 1 female, ex pupa, found on Guava (Psidium guajava L.), A. & M. E. Blanchard 
coll. Same locality, 3 larvae, 2 pupae collected on Guava by R. O. Kendall, producing 
the following adults: 24-XI-69, 3 females; 23-XII-69, 1 female. 1 larva preserved. Det. 
E. Todd and also recorded from Texas by him as follows: Brownsville, Oct. 1953, J. M. 
McGough coll.; Mercedes, July 1957, P. T. Riherd coll. Ground color light gray; post- 
medial line bordered inwardly with orange, outwardly with blackish triangular spots; 
pattern otherwise black. 


Geometridae 


Scopula eburneata Guenée (Fig. 17). All from North Padre Island, either Kleberg or 
Nueces Co., 17-V-76, 2 females; 24-VI-76, 1 male; 19-VII-76, 1 male; 19-VI-77, 1 male; 
21-VI-77, 1 female; 6-IV-78, 2 females; 8-11-VI-78, 1 male, 3 females, all A. & M. E. 
Blanchard coll.; 1-X-77, 2 males, E. Knudson coll. Det. D. S. Fletcher (BM). Ground 
color creamy white with blackish markings. 


Pyralidae 


Araschnopsis subulalis (Guenée) (Fig. 18). Hidalgo Co., Bentsen Rio Grande Valley 
St. Pk., 20-X-74, 1 female, E. Knudson coll.; Santa Ana Refuge, 28-XI-75, 1 female, A. 
& M. E. Blanchard coll. New U.S. record. Det. E. G. Munroe and D. C. Ferguson. 
Creamy white reticulated with medium brown; brown patches on outer margins of both 
wings. 

Lamprosema baracoalis Schaus (Fig. 19). Hidalgo Co., Santa Ana Refuge, 5-III-73, 1 
female; 28-XI-75, 1 female, A. & M. E. Blanchard coll.; same locality, 20-X-74, 1 female, 
E. Knudson coll. Det. D. C. Ferguson. Chocolate-brown with hyaline spots on forewing 
(basal area in figure partially denuded of scales by wear); hindwing with narrow white 
antemedial line, bordered inwardly with black. 

Bocchoropsis pharaxalis Druce (Fig. 20). Hidalgo Co., Santa Ana Refuge, 25-XI-73, 1 
female, A. & M. E. Blanchard coll. New U.S. record. Det. D. C. Ferguson. Shiny yel- 
lowish white marked with light brown. 

Stemhorrhages costata (Fab.) (Fig. 21). Harris Co., Bellaire, 15-IX-80, 1 male; 25-IX- 
83, 1 male; Kerr Co., Hunt, 23-VIII-81, 1 female, E. Knudson coll.; Dallas Co., Garland, 
14-IX-83, 1 female, H. A. Freeman coll. Det. D. C. Ferguson. This species is also reported 


— 


Fics. 8-14. 8, Letis xylia (Guenée), Padre Island Nat’l. Seashore, 19-VII-76, wing 
expanse 82 mm. 9, Cropia ruthaea Dyar, Brewster Co., Big Bend Nat’l. Park, Green 


VOLUME 39, NUMBER 1 


Gulch, 28-V-81, wing expanse 35.9 mm. 10, Hemispragueia idella Barnes, Presidio Co., 
Ruidosa Hot Springs, 8-VII-69, wing expanse 23.5 mm. 11, Lesmone fufius (Schaus), 
Hidalgo Co., Santa Ana Refuge, 13-IX-80, wing expanse 26.2 mm. 12, Lesmone for- 
mularis (Geyer), Hidalgo Co., Santa Ana Refuge, 28-XI-75, wing expanse 30.8 mm. 13, 
Eulepidotis addens (Walker), Hidalgo Co., Bentsen Rio Grande Valley St. Pk., 11-X-80, 
wing expanse 26.8 mm. 14, Gonocarsia electrica Schaus, Brewster Co., Big Bend Nat’. 
Park, Green Gulch, 7-X-66, wing expanse 40.5 mm. (Figures not at same scale. Full wing 
expanse given for each specimen. ) 





6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


from Louisiana for the first time by V. A. Brou as follows: West Feliciana Par., Weyanoke, 
30-IX-79, 1 female; St. John Par., Edgard, 12-V-82, 1 male; 6-I[X-82, 1 female, V. A. 
Brou coll. Glossy white with yellow-brown costa and small black discal spot on forewing; 
male with prominent black anal tuft. 

Daulia arizonensis Munroe (Fig. 22). Brewster Co., Big Bend Nat'l. Park, Chihuahuan 
desert near Nugent Mt., 21-IX-71, 2 males; Gov't. Spring, 29-IX-65, 1 female; 11-X-69, 
1 male, A. & M. E. Blanchard coll. Big Bend Nat’l. Park, Gov't. Spring, 28-IX-81, 1 male; 
K-Bar Research Station, 28-IJJ-83, 1 male; Study Butte, 4-XJ-81, 2 males, 1 female; El 
Paso Co., Tom Mays Park, 24-V-81, 1 male, 1 female, E. Knudson coll. Det. E. G. Munroe. 
Forewing golden yellow with silvery markings, margined with black scales; hindwing 
whitish. 

Microthyris prolongalis Guenée (Fig. 23). Hidalgo Co., Santa Ana Refuge, 28-XI-75, 
1 male, A. & M. E. Blanchard coll. Det. D. C. Ferguson. Medium brown with obscure 
hyaline spots on forewing; hindwing with dark brown antemedial line. 

Salobrena sincera (Zeller) (Fig. 24). A common and widely distributed species in 
southwestern Texas, from San Antonio south to Brownsville and west to the trans-Pecos 
region. Varying shades of brown tinted with reddish orange; forewing of male with huge 
funnel-shaped costal process; female with much smaller costal process. This common 
Texas species is included to illustrate the peculiar wing shape of the male. 

Hemiplatytes parallela (Kft.) (Fig. 25). Randall Co., Palo Duro Canyon, 11-IX-66, 1 
male, A. & M. E. Blanchard coll.; same locality, 9-V-81, 2 females; Brewster Co., Alpine, 
5-V-78, 1 male, E. Knudson coll. Det. D. C. Ferguson. Forewing light yellowish brown 
with broad, continuous silvery streak from base to apex, and narrow silvery streak on 
fold; hindwing white. 

Diviana eudoreella Ragonot (Fig. 26). Kimble Co., Junction, 16-IV-74, 1 male, A. & 
M. E. Blanchard coll.; Jeff Davis Co., Davis Mountains St. Pk., 28-V-79, 1 female; Mt. 
Locke, 8-VIII-83, 1 female; Kerr Co., Hunt, 4-X-80, 1 male (donated to USNM), 1 female; 
25-VII-81, 1 male; 6-IX-81, 1 male, all E. Knudson coll. Det. D. C. Ferguson. Forewing 
light gray with blackish gray markings; hindwing pale fuscous. 

Sosipatra nonparilella Dyar (Fig. 27). Jeff Davis Co., Ft. Davis, 13-X-66, 1 female; 
Brewster Co., Big Bend Nat’l. Park, Chihuahuan desert near Nugent Mt., 1-V-72, 1 male, 
A. & M. E. Blanchard coll. Forewings whitish gray, evenly speckled with black scales; 
lines white, nearly straight, bordered with black; hindwing pale fuscous. This rare species 
is similar in appearance to several other phycitines, but may be separated by genitalic 
characters. 


Tortricidae 


Sparganothis cana Rob. (Fig. 28). Walker Co., Huntsville St. Pk., 20-V-69, 5 males, 1 
female, A. & M. E. Blanchard coll. Det. J. Powell. Forewings light gray, reticulated with 
black; the two costal and one dorsal patches are medium gray; hindwing fuscous. 


Cossidae 


Hypopta palmata B. & McD. (Fig. 16). Brewster Co., Big Bend Nat’l. Park, Green 
Gulch, 29-IX-81, 1 male; Dugout Wells, 9-VIII-83, 1 male, E. Knudson coll. Forewing 


~ 


Fics. 15-28. 15, Nystalea collaris (Schaus), Cameron Co., Brownsville, 8-XI-69, 
wing expanse 51 mm. 16, Hypopta palmata B. & McD., Brewster Co., Big Bend Nat'l. 
Park, Green Gulch, 29-IX-81, wing expanse 30 mm. 17, Scopula eburneata Guenée, 
North Padre Island, 19-VI-77, wing expanse 13 mm. 18, Araschnopsis subulalis (Gue- 
née), Hidalgo Co., Santa Ana Refuge, 28-XI-75, wing expanse 20 mm. 19, Lamprosema 
baracoalis Schaus, Hidalgo Co., Santa Ana Refuge, 28-XI-75, wing expanse 22.3 mm. 
20, Bocchoropsis pharaxalis Druce, Hidalgo Co., Santa Ana Refuge, 25-XI-73, wing 


VOLUME 39, NUMBER 1 7 





expanse 20.3 mm. 21, Stemhorrhages costata (Fab.), Harris Co., Bellaire, 15-IX-80, wing 
expanse 29.6 mm. 22, Daulia arizonensis Munroe, Brewster Co., Big Bend Nat'l. Park, 
Chihuahuan desert near Nugent Mt., 21-IX-7], wing expanse 29.6 mm. 23, Microthyris 
prolongalis Guenée, Hidalgo Co., Santa Ana Refuge, 28-XI-75, wing expanse 29.6 mm. 
24, Salobrena sincera (Zeller), Cameron Co., Brownsville, 12-IV-66, male. wing expanse 
13.6 mm. 25, Hemiplatytes parallela (Kearfott), Randall Co., Palo Duro Canyon, 11- 
IX-66, wing expanse 25.1 mm. 26, Diviana eudoreella Ragonot, Kimble Co., Junction, 
16-IV-74, wing expanse 15.8 mm. 27, Sosipatra nonparilella Dyar, Jeff Davis Co., Ft. 
Davis, 13-X-66, wing expanse 20.1 mm. 28, Sparganothis cana Robinson, Walker Co.., 
Huntsville St. Pk., 20-V-69, wing expanse 18.7 mm. (Figures not at same scale. Full wing 
expanse given for each specimen. ) 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


whitish, irrorated with black scales; veins black; dark gray patches on fold and subter- 
minal area; hindwing fuscous. 


ACKNOWLEDGMENTS 


The authors are extremely grateful for the assistance of Drs. D. C. Ferguson, D. S. 
Fletcher, J. G. Franclemont, R. W. Hodges, J. D. Lafontaine, E. G. Munroe, R. B. Poole, 
J. A. Powell, and E. L. Todd for determinations and especially to Drs. Ferguson and 
Poole for reviewing the manuscript. We are also grateful to R. O. Kendall, V. A. Brou, 
and Dr. E. L. Todd for providing additional records. We also wish to express thanks to 
the Texas Parks and Wildlife Dept. and U.S. National Park Service for their assistance. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 9-12 


NOTES ON THE LARVA AND BIOLOGY OF 
MOODNA BISINUELLA HAMPSON 
(PYRALIDAE: PHYCITINAE)! 


H. H. NEUNZIG 


Department of Entomology, North Carolina State University, 
Raleigh, North Carolina 27650 


ABSTRACT. The last stage larva of Moodna bisinuella Hampson is described, and 
the biology of this phycitine with reference to gama grass (Tripsacum) in North Carolina 
is briefly outlined. 


Recently, gama grass (Tripsacum sp.) was brought to Raleigh, North 
Carolina from Mexico and planted on a research farm as part of a 
plant breeding program. Inadvertently, phycitine larvae were intro- 
duced with the plants. Injury to the plants eventually prompted the 
collection of larvae and rearing of adults. Adults were identified as 
Moodna bisinuella Hampson, a species of economic importance in 
Central America. Although the phycitines were eradicated from the 
grass in North Carolina after the insects were identified, notes taken 
and larval specimens obtained during the rearing procedures provided 
worthwhile information relative to this pest. Little has been published 
regarding the appearance of the immature stages and biology of M. 
bisinuella. Previous authors have only mentioned a few morphological 
features of the larva (Capps, 1963) and merely stated that the species 
feeds as a larva in the ears of soft or “green” corn (Zea mays L.) 
(Capps, 1968; Heinrich, 1956). In this paper, I describe the last stage 
larva in detail and briefly discuss the biology of M. bisinuella in as- 
sociation with its previously unreported host, gama grass. 


Description of Last Stage Larva 


General. Length 10.2-16.0 mm, avg. 13.5 mm. 

Color. Head yellowish brown (at times with green undertones in living larva); tono- 
fibrillary platelets pale brown, indistinct; usually, a pale brown to brown patch within 
arc of stemmata and pale brown to brown streaks near notch of postgenal region (these 
2 pigmented areas sometimes coalescing); hypostoma with brown to black markings; 
mandibles yellowish brown between articulations becoming dark brown along lateral 
margins and distally. 

Prothoracic shield pale yellow to pale brown with lateral and posterior margins darker 
(green undertones in living larva). Prespiracular plate yellowish brown to dark brown. 
Remainder of prothorax white to yellowish white overlaid with brown to gray integu- 


‘Paper No. 9067 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, North Carolina 
27650. 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1 2 


Fics. 1, 2. Moodna bisinuella Hampson. 1, mesal aspect of right mandible of last 
stage larva. 2, dorsal aspect of part of right maxilla of last stage larva. 


mental granules (living larva with remainder of prothorax pale red to red with brown 
to gray granules and scattered blue undertones; red pigmentation usually more intense 
on mesothorax and metathorax, and blue more pronounced laterally and ventrally); 
mesothoracic pinaculum ring pale brown to dark brown, white within ring; thoracic legs 
mostly pale brown to brown. 

Abdomen similar to mesothorax and metathorax; eighth abdominal segment pinacu- 
lum ring pale brown to brown; anal shield pale yellow to yellowish brown with slightly 
darker margins. 

Morphological features: Head. Width 0.83-0.93 mm, avg. 0.90 mm; surface slightly 
uneven; adfrontals reach ca. % distance to epicranial notch; AF2 setae usually slightly 
ventrad of level at which epicranial suture forks; AF2 setae slightly above imaginary line 
between P1 setae; P1 setae further apart than P2 setae; labrum shallowly notched; man- 
dibles simple, distal teeth distinct (Fig. 1); sensilla trichodea of maxillae simple (Fig. 2). 
Spinneret moderately long. 

Prothorax. On shield, distance between D1 setae less than distance between XD1 
setae; on each side of shield, distance between SD1 and SD2 greater than distance 
between SD1 and XD2, distance between D1 and D2 greater than distance between D1 
and XD1, and XD2, SD1 and SD2 form an acute angle; L setae of each side in a nearly 
vertical configuration. 

Mesothorax and metathorax. SD1 pinaculum rings of mesothorax well developed; 
SD1 setae of mesothorax ca. 2 times as long as SD1 setae of metathorax; SD1 and SD2 
pinacula of metathorax fused; D1 and D2 pinacula of metathorax fused. 

Abdomen. D2 setae of anterior segments ca. 0.4 mm long; D1 setae of anterior seg- 
ments ca. as long as D2 setae; distance between D2 setae on segments 1-7 slightly greater 
than distance between D1 setae; distance between D1 and D2 on each side of segments 
3-6 about same as distance between D1 and SD1; SD1 setae of segments 1-7 without 
pinaculum rings; crochets in a biordinal ellipse, avg. number on prolegs of segments 3, 
4, 5, 6, and anal segment, 50, 56, 57, 58, and 49, respectively; vertical diam. of spiracles 
on segment 8 ca. % larger than same diam. of spiracles on segment 7; horizontal diameter 
of spiracle of each side of segment 8 slightly less than distance between L1 and L2; SD1 
pinaculum rings of segment 8 relatively broad but appearing incomplete; SD1 setae of 
segment 8 ca. 1.9 times longer than SD1 setae of segment 7; 2 SV setae on each side of 
segment 8; distance between D1 and D2 on each side of segment 9 ca. 3 times distance 
between D1 and SD1; 2 SV setae on each side of segment 9. 


VOLUME 39, NUMBER 1 ie 





Fics. 8, 4. Seeds of gama grass infested with last stage larvae of Moodna bisinuella 
Hampson. 


Material Examined 


North Carolina: Raleigh; 5 larvae, Tripsacum sp. seed, 19-IX-80, 
Coll. H. H. Neunzig; 11 larvae, Tripsacum sp. seed, 30-IX-80, Coll. H. 
H. Neunzig. These specimens have been deposited in the NCSU Insect 
Collection. 


BIOLOGY AND DISCUSSION 


In North Carolina, M. bisinuella overwintered as diapausing larvae 
within silk enclosures constructed at the inner base of gama grass. 
Pupation occurred in the spring, and adults emerged in April and May. 
Oviposition and larval feeding sites for the spring generation could not 
be determined. Larvae of the summer generation (July-September) 
were all found associated with the seeds of gama grass. These larvae 
fed on the well-developed, but more or less soft, seeds. Usually this 
occurred while the seeds were still attached to the plant, but seeds that 
had fallen from the plant were also eaten. The seeds were bored into 
and frass and silk extruded from the entrance hole (Figs. 3 & 4). Several 
seeds were eaten by each larva as it developed. Ergot (Claviceps sp.), 
which was at times associated with gama grass seed, was also sometimes 
ingested by the larvae. 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The fact that M. bisinuella feeds on gama grass as well as corn 
supports the botanical view that the two plant genera (Tripsacum and 
Zea) are closely related (they are considered by most botanists to be 
the only members of the New World tribe Maydeae). In Central Amer- 
ica, where the two plants sometimes grow in close proximity, wild 
communities of gama grass are in all likelihood providing a reservoir 
of M. bisinuella that periodically infest fields of corn. 


ACKNOWLEDGMENTS 


D. L. Stephan of the Plant Disease and Insect Clinic of North Carolina State University 
made available to the author the initial series of larvae of M. bisinuella collected in 
North Carolina. 


LITERATURE CITED 


Capps, H. W. 1963. Keys for the identification of some lepidopterous larvae frequently 
intercepted at quarantines. U.S. Dept. Agr. ARS-33-30-1. 37 pp. 

HEINRICH, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus. Bull. 
207. 581 pp. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 13-18 


BIOLOGY AND DESCRIPTION OF THE LARVA OF 
DICYMOLOMIA METALLIFERALIS: A CASE-BEARING 
GLAPHYRIINE (PYRALIDAE) 


DAVID WAGNER 


Department of Entomology, 218 Wellman Hall, 
University of California, Berkeley, California 94720 


ABSTRACT. The larval biology of Dicymolomia metalliferalis (Packard) is de- 
scribed. Larvae were collected from partially opened, necrotic seed pods of two Lupinus 
L. species and reared to adults. Larvae fed from within purse-like cases constructed of 
silk and detrital tissues. This constitutes the first record of the case-bearing habit for the 
glaphyriine genus, Dicymolomia Zeller. The larva is described and compared with that 
of D. julianalis (Walker). 


Dicymolomia metalliferalis (Packard) is a day-flying pyralid found 
from southern Vancouver Island south to San Diego (Munroe, 1972). 
Although adults (Fig. 1) may be locally common, the early stages of 
this moth were previously unknown. During the course of a faunal 
survey of the Lepidoptera associated with Lupinus L. species, I reared 
D. metalliferalis from partially opened, decaying seed pods of two 
perennial Lupinus species. 

Larvae were recovered from the seed pods of the previous spring’s 
seed set. In most of the inflorescences surveyed, greater than 95% of 
the seed pods had fully dehisced, releasing all their seeds. However, a 
fraction of the seed pods had failed to open completely; these partially 
intact pods were split open and examined for lepidopteran larvae. Over 
400 seed pods of Lupinus arboreus Sims (ca. 150), L. chamissonis 
Eschs. (ca. 80), L. albifrons Benth. (ca. 150), and L. latifolius J. G. 
Agardh (ca. 20) were examined; D. metalliferalis larvae were re- 
covered from the latter two species: CA, Marin Co., Nicasio Reservoir, 
26-I-1980, ex L. albifrons; Contra Costa Co., Tilden Park, nr. Inspi- 
ration Point, 2-IJ-1980, ex L. latifolius; and Contra Costa Co., Briones 
Reservoir, 17-I-1982 and 12-II-1983, ex L. albifrons. 

The partially intact seed pods from which larvae were recovered 
invariably showed signs of insect feeding damage. Many of the occu- 
pied pods were partially held together by the silk of braconid cocoons 
which presumably resulted from the parasitism of seed-feeding Lepi- 
doptera. Other seed pods hosting Dicymolomia larvae had been at- 
tacked by Apion Herbst, a common seed-feeding curculionid that oc- 
curred in all Lupinus species studied. No larvae were collected from 
intact seed pods; presumably, Dicymolomia larvae entered damaged 
seed pods secondarily. 

Most of the larvae were collected in purse-like cases (Fig. 2), but 
several larvae were recovered from chambers of sparse silk formed in 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





PETEETET EET 
Fics. 1 & 2. 1, Dicymolomia metalliferalis, Briones Reservoir, Contra Costa Co., 
CA. 2, early instar cases, lower right case opened. Scale in mm. 


cavities of the partially intact fruits. However, soon after being brought 
indoors, these larvae constructed cases. After one week all larvae were 
observed feeding from within cases where they remained to maturity. 
Larvae were reared on necrotic seed pod tissues or on a combination 
of pod fragments and wheat germ. 

Cases were constructed of silk with miscellaneous inclusions. One 
field-collected, overwintering case (length: 0.48 mm) contained por- 
tions of a seed coat, seed pod tomentum, larval frass, and an earlier 
instar head capsule. The case of a mature laboratory-reared larva 
(length: 1.3 cm) incituded numerous leaf fragments and wheat germ. 
Several of the field-collected cases had at least two species of fungi 
growing on them, and associated mites which presumably were feeding 
on the fungi. Neither the fungi nor the mites appeared to have had a 
detrimental effect on the larvae. Larval cases were constructed with 
an opening at each end. 

Pupation occurred within the larval cases. Laboratory-reared adults 
emerged between 15 May and 25 June (N = 3). However, most of the 
records for field-collected adults occur later in the summer: CA, Berke- 
ley, 23-VI to 10-IX (12 records, CIS collection). 

The larval description is based on six late instar larvae collected at 
Briones Reservoir, Contra Costa Co., CA, on 17-I-1982 (1) and 12-II- 
1983 (5). Larvae were distended in hot water and then transferred to 
70% EtOH. One larva was cleared in 10% KOH and stained with 
chlorosol black to facilitate the examination of smaller setae. Measure- 
ments refer to a mature fully distended larva. Setal nomenclature fol- 
lows Hinton (1946). 


DISCUSSION 


Six species of Dicymolomia are reported for North America north 
of Mexico (Munroe, 1972). Larvae of the genus have not been de- 


VOLUME 39, NUMBER 1 15 


scribed or figured in detail. However, Forbes (1932) included a setal 
map of D. julianalis (Walker) for segments T1, T2, A2, and A7-9. 
Larvae of D. metalliferalis are readily separable from those of D. 
julianalis. The mesothorax of D. julianalis has a large sclerotized shield 
extending from just anterodorsad of D1 to the dorsal midline, which 
is absent in D. metalliferalis. In D. metalliferalis L1 and L2 on T2- 
A8 and the SV setae on A2-7 are not included on the same pinaculum 
as in D. julianalis. Forbes did not illustrate an SD2 seta in D. julianalis, 
which is present but minute in D. metalliferalis. 

D. julianalis exhibits a diverse range of larval substrates: larvae are 
recorded from Astragalus canadensis L. and Cirsium lecontei Torrey 
and Gray; as internal feeders in cat-tails (Typhus) and cactus stems 
(Opuntia); from senescent cotton bolls (Gossypium); and as predators 
on the eggs and larvae of bagworms, Thyridopteryx ephemeraeformis 
(Haworth) (Munroe, 1972). Although D. metalliferalis larvae have been 
reared from only two lupine species, circumstantial evidence suggests 
that this moth may utilize an array of larval substrates. Larvae were 
recovered in rather low densities relative to numbers of adults flying 
at the collection sites. Furthermore, D. metalliferalis adults may be 
collected in localities with little or no lupine. The decomposed nature 
of the occupied pods suggests that larvae feed generally on detrital 
tissues. Lastly, one overwintering larva was reared to maturity on a 
mixture of seed pod fragments and wheat germ. 

Two other species of microlepidoptera were commonly associated 
with partially intact, necrotic Lupinus seed pods. Several fully grown 
overwintering larvae of Argyrotaenia citrana (Fern.) (Tortricidae) were 
collected inside seed pods; larvae pupated soon after their collection 
with no indication as to having fed on the seed pods. A member of an 
undetermined gelechiine genus was frequently collected in the seed 
pods of L. arboreus, L. albifrons, and L. latifolius. Laboratory-reared 
larvae were observed to feed on the necrotic tissues of the fruit pericarp 
beneath sheets of silk. The larvae were smokey-red in color with pale 
longitudinal stripes. 

The case-bearing habit is of general occurrence in several primitive 
lepidopteran taxa: in the later instars of Incurvarioidea, e.g., Adelinae 
and Incurvariinae; in the Tinoidea, e.g., Tineidae and Psychidae; and 
in the Gelechioidea, e.g., Coleophoridae and some Oeccophoridae 
(Common, 1970). Case-bearing is of sporadic occurrence in the Tor- 
tricoidea, e.g., Clysiana acrographa (Turn.) (Common, 1970) and Pyr- 
aloidea, e.g., Nymphulinae and Glaphyriinae. Within the Glaphyri- 
inae, case-bearing has been reported for two genera: Stegea Munroe 
and Lipocosma Lederer (Forbes, 1932; Munroe, 1972). Hence, Dicy- 
molomia is the third of 15 North American glaphyriine genera for 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





6mm 


Fics. 3-8. Head of D. metalliferalis. 3, head capsule, dorsal view, width 0.93 mm. 
4, head capsule, lateral view. 5, labrum. 6, antenna. 7, mandible, mesal view. 8, maxilla. 


which case-bearing is known. Detrivory appears to be especially com- 
mon among case-bearing taxa relative to other Lepidoptera, e.g., at 
least some Adelinae, Tineidae, Oecophoridae, and Dicymolomia. 


DESCRIPTION OF MATURE LARVA 


General. Overall length 10.2 mm. Body salmon-orange, intersegmental regions unpig- 
mented in living larvae, color fading to white in preserved material. Setulae indistinct, 


VOLUME 39, NUMBER 1 17 





meson 


Fics. 9-11. Thorax and abdomen of D. metalliferalis. 9, setal map of segments T1- 
2, Al-3, & A7-10. 10, dorsal view of A8—10. 11, crochets on A6. 


visible at 100. Pinacula indistinct. Spiracles flat, lightly pigmented; spiracles on T1 and 
A8 twice the diameter of those on Al-7. 

Head. (Figs. 3, 4) 0.93 mm wide; frons extending “% of the way to occipital foramen. 
Head capsule yellow-brown to red-brown. Six stemmata; $1, $2, $3, S4 and S6 equidistant, 
forming a semicircle; S5 below antennae; S1 almost twice the diameter of other stemmata. 
Labrum (Fig. 5) 1.25x as broad as long; anterior margin emarginate and heavily pig- 
mented. Antenna (Fig. 6) 0.15 mm; scape lightly pigmented; scape and pedicel subequal 
in length. Mandible (Fig. 7) with five teeth, ventral tooth notched; molar process mod- 
erately developed. Maxilla (Fig. 8) 0.42 mm; stipes with an elongate sclerite along mesal 


18 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


margin and small free sclerite at level of the spinneret; palpiger and palpus lightly 
sclerotized. Labium with a hook-like sclerite on either side of spinneret. 

Thorax. Prothoracic shield brown; L1 and L2 on a darkened pinaculum; SV setae on 
lightly sclerotized pinaculum; 5 coxal setae. Meso- and metathorax with sclerite poster- 
iorad of D and SD setae, smaller and paler on metathorax. 

Abdomen. (Figs. 9, 10) D1 twice the length of D2 on A1-7; subequal on A8-9. SD1 
large, directly above spiracle; SD2 minute. L1 and L2 approximate and in vertical row. 
Al with two and A2 with three SV setae. A9 with a pair of subcuticular pigment spots 
near dorsal midline. Al0 with 13 pairs of setae; anal plate irregular with numerous 
subcuticular pigment spots laterad of anal plate (Fig. 10). Crochets 16-28 in biordinal 
penellipse on A3-6 (Fig. 11), anterior crochets slightly larger; crochets 15-26 in a bior- 
dinal transverse band on A10. 


ACKNOWLEDGMENTS 


I thank Jerry A. Powell and John A. DeBenedictis for their helpful suggestions and 
reviewing drafts of this paper. 


LITERATURE CITED 


ComMoON, I. F. B. 1970. In The insects of Australia. Melbourne Univ. Press, Canberra. 
802 pp. 

FORBES, W. T. M. 1932. The Lepidoptera of New York and neighboring states. Cornell 
Univ. Press, Ithaca. 536 pp. 

HINTON, H. E. 1946. On the homology and nomenclature of the setae of the lepidop- 
terous larvae, with some notes on the phylogeny of the Lepidoptera. Trans. R. Ento- 
mol. Soc. Lond. 97:1-87. 

MUNROE, E. E. 1972. In Dominick et al. The moths of North America north of Mexico. 
Fasc. 13.1B. Pyraloidea. E. W. Classey Ltd. & R. B. D. Pub. Inc., London. Pp. 194— 
250. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 19-25 


FIELD SURVEY OF THE TRUE BUTTERFLIES 
(PAPILIONOIDEA) OF RHODE ISLAND 


HARRY PAVULAAN 
1919 North Daniel St., #201, Arlington, Virginia 22201 


ABSTRACT. The survey was undertaken to establish a better understanding of but- 
terfly occurrences in an area previously lacking in published records for many species of 
the Papilionoidea. All species observed in the field are indigenous to the entire New 
England region, although some are very selective in choosing their particular habitats. 
It is interesting to note that many of the species listed are technically records for the 
state. 


Following are the results of my 1988 field studies of the butterflies 
of Rhode Island. Most occurrences were confirmed by several captures 
at that particular location (usually followed by release of the specimen), 
but many sightings were logged on the basis of behavioral character- 
istics. This task was especially easy in the case of the most common 
species, while the rarer species were recorded only after a documented 
capture. Doubtful or questionable sightings were not recorded. 

Most sightings were recorded from five primary study areas (Fig. 1) 
which were visited several times per week on a regular basis through- 
out the collecting season: 

1. Trestle Trail, Coventry town: A dismantled railroad bed offering 
an excellent cross-section of the region’s predominant oak forest with 
several areas of woodland swamp. This proved to be a poor area for 
butterflies, containing a fair variety of species but generally in very 
low numbers, totalling 17. Several oak tree species and an undergrowth 
of various blueberries predominate here, with wildflowers generally 
lacking. 

2. West Warwick and eastern Coventry towns: Old established res- 
idential/commercial/industrial area interspersed with urban lots, 
abandoned areas, disturbed fields, and neglected weedy areas. Wild- 
flowers such as asters, daisies, goldenrod, milkweed, parsnip, sunflow- 
ers, and loosestrife (in wet areas) abound in these open places and 
attract an abundance of the more typical weed-field butterfly species, 
totalling 23. 

3. Arcadia Wildlife Management Area, Exeter: Extensive areas of 
oak and pitch pine forest traversed by a network of dirt roads. The 
area contains several large weed fields. A fair variety of species occurs 
here, totalling 16; but again, numbers are very low. 

4. Great Swamp Wildlife Management Area, South Kingstown: 
Mostly oak forest with areas of mixed forest, extensive freshwater wet- 
land, and several large weed fields. Other unique features such as 


20 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





PROVIDENCE 





BRISTOL 





WASHINGTON 


9 NEWPORT 


Y Primary Study Area 


@ Secondary Study Area 


Win Route 2 Corridor 


Fic. 1. Study areas for field survey of butterflies of Rhode Island. 


sphagnum and cranberry bogs, maple swamp, and at least two stands 
of Atlantic white cedar are found here. This is perhaps the richest 
ecological area of the state, containing the largest variety of wildflowers 
and butterflies, which total 26 species. Many species reach their great- 
est abundance here. 

5. East Matunuck State Beach, South Kingstown: A typical east 
coast beach, with predominant dune grass, rugosa rose thickets, and 
late-season goldenrods. Not a very rich area for butterflies, except to- 
ward the season’s end, at which time butterfly numbers seem to in- 


VOLUME 39, NUMBER 1 Pa). 


crease remarkably. Probably due to the ocean’s moderating effect on 
early frosts experienced inland, one will find butterflies here weeks 
after they have disappeared in all other locations. Only 10 species were 
recorded here. 

Several secondary study areas were also included in this survey (Fig. 
1). These were each visited on an irregular basis although a very thor- 
ough count was made on each occasion: 

6. George Washington Wildlife Management Area, Burrillville: 
Mixed transition zone forest interspersed with wooded swamps and 
lakes. A surprisingly poor area for butterflies, with only 3 species sight- 
ed. Visited in early August. 

7. Durfee Hill Wildlife Management Area, Glocester: An open wet- 
land bordered by mixed forest and a field of goldenrod and joe-pye 
weed. Colias species abundant but few others are present. Only 5 
species were sighted in late August. 

8. Trustom Pond Wildlife Refuge, South Kingstown: Characterized 
by coastal mixed forest with abundant scrub oak. There are several 
large weed fields here. 15 species of butterflies were recorded here in 
late July. 

9. Beavertail State Park, Conanicut Island, Jamestown: The weed 
fields above this rocky coastal hillside are characterized by thorny scrub 
and an abundance of goldenrods and other wildflowers. Pierids abun- 
dant with some noticeable gathering of fall migratory species, totalling 
19. Visited throughout September. 

10. Ninigret Conservation Area, Charlestown: Typical east coast 
barrier beach. Same plants and conditions as in area 5. However, only 
6 species were sighted here from late September to early October. 

Finally, sightings were recorded along local roads, and generally 
anywhere, when possible, during my various treks throughout the state: 

11. State Route 2, Warwick City to South Kingstown: Cutting a 
cross-sectional corridor through the state’s heartland, this was the most 
thoroughly logged road in my study. Observations were made at var- 
ious roadside stops and at random locations off the route. This road 
bisects a variety of environments, from suburban development to 
woodlands, weedy fields, and agriculture. Observations indicate that 
butterflies are generally scarce except common pierids. Only 6 species 
were recorded throughout the year. 

12. Various other places throughout the state, wherever I travelled, 
were subject to investigation. None were ever investigated to any de- 
gree beyond, perhaps, a one hour collecting stop. A total of 18 species 
was recorded outside the 11 study areas. 

1983 was characterized by abnormal weather patterns. Although 
blessed by one of the mildest winters on record, the spring was unusu- 


22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ally cool, cloudy, and brought record rainfalls to New England. This 
had detrimental effects on spring broods, keeping numbers low. The 
summer was characterized by an extended period of drought and ex- 
cessive heat, lasting well into September. This probably also spelled ill 
for summer broods, keeping numbers low except in the case of the 
common weed-field species which seemed to flourish. The fall started 
off mildly, but cold weather set in suddenly in mid-October, with some 
frosts inland. 

In general, collecting was very poor in 1983 in Rhode Island, and 
apparently much of New England as a whole. The most common 
species appeared in healthy numbers, but less common species and 
“exotics” were found to be either very low in numbers, rare, or absent 
altogether. 


RESULTS 


Following each description, numbers refer to all study areas (Fig. 1) 
in which sightings were made, and in parentheses are county listings, 
abbreviated as follows: B = Bristol, K = Kent, N = Newport, P = Prov- 
idence, W = Washington. New county records are in italics. State rec- 
ords are denoted by an asterisk (*). 


Danaus plexippus plexipnpus Linnaeus: Northward migrants first appeared in moderate 
numbers in mid-vii but became scarce thereafter. Fresh adults appeared in small numbers 
in milkweed fields during late-viii, but dispersed widely by 1-ix. Population swelled in 
the southern part of the state from mid-ix to early-x, with gathering along the coast but 
no obvious migratory movement. Areas 1, 2, 4, 5, 7, 8, 9, 10, 11, 12. (KNPW). 

Celastrina laden ladon Cramer: Appeared in great abundance in most localities in iv. 
Usually found along dirt roads in wooded areas. Form lucia dominant early, marginata 
later, and violacea dominant toward end of brood in v, with all degrees of intergrades 
present throughout. I have assembled a complete series of these intergrades, ranging 
from very heavily-marked lucia, through marginata, to very light, sharply-marked vio- 
lacea. In contrast, summer form neglecta was represented by only a few isolated sightings 
in vii. I suspect a possible sibling species relationship here. Spring forms lucia, marginata, 
violacea in areas 1, 2, 3, 4, 11, 12. (KW).* Summer form neglecta in areas 1, 2, 12. 
(KP).* 

Celastrina neglecta-major Tutt (?): Found mainly at Great Swamp. Originally thought 
to be a partial second brood of C. ladon, but evidence suggests that this butterfly is, very 
likely, a sibling species. Appears toward the end of C. ladon spring brood in v, when 
large, bright blue, freshly-emerged males of neglecta-major mingle with worn stragglers 
of C. ladon form violacea. The neglecta-major males seemed to disregard C. ladon 
females with which they flew, apparently seeking their own kind. The females are very 
elusive, keeping to the woods and avoiding open spaces. One female was observed flying 
about a dogwood tree at Great Swamp. Areas 4, 12. (KW).* 

Everes comyntas comyntas Godart: Three broods. A small brood in early-vi, as the 
spring form, again appearing in small numbers in early-vii as the summer form. Common 
only in viii as a full summer brood, with late emergers into early-x. Found mostly in 
weed fields. Areas 2, 3, 4, 7, 12. (KPN).* 

Incisalia henrici henrici Grote & Robinson: Appearing in small numbers in early-v in 
the vicinity of Great Swamp. Found only in association with American holly, never 


VOLUME 39, NUMBER 1 23 


straying far from this plant, although another reported foodplant, highbush blueberry, 
is abundant here. Area 4. (W).* 

Incisalia niphon clarki Freeman: Represented by one individual captured in early-v 
in the vicinity of Great Swamp. Found in close proximity to a stand of pitch pine, the 
reported foodplant. Area 4. (W).* 

Lycaena phlaeas americana Harris: Three broods. Spring form common from late-v 
to early-vi. Summer form in two broods of moderate numbers, throughout July and again 
in late-viii through ix. Found mostly in weed fields. Areas 1, 2, 3, 4, 8, 9. (KWN).* 

Satyrium acadica acadica Edwards: Represented by one individual taken in mid-vii 
in West Warwick. This butterfly was sighted near a pond ringed with willows, the 
reported foodplant. Area 2. (K).* 

Satyrium edwardsii Grote & Robinson: Very few of these butterflies were taken in 
oak woods in mid-vii. One sighting near the coast in a scrub oak thicket. Areas 1, 8. 
(KW). 

Satyrium liparops strigosus Harris: Very few captures in oak woods, from late-vii to 
early-viii. Areas 1, 2, 3. (KW). 

Strymon melinus humuli Harris: Surprisingly only one sighting of this butterfly, which 
is normally common in the northeast. Captured in West Warwick, late-vii. Area 2. (K). 

Strymon titus titus Fabricius: Very few captures in oak woods throughout vii. Areas 
1, 2, 4. (KW). 

Boloria selene myrinna Cramer: Only two individuals of this reputedly common species 
were sighted at Great Swamp in mid-vii. Area 4. (W). 

Limenitis archippus archippus Cramer: The first brood in v produced very few in- 
dividuals. An extended summer brood produced individuals in two overlapping cycles, 
possibly a “split” second brood. Males appeared in moderate numbers at most localities 
in mid-vii, with females first becoming evident only one week later. Most individuals 
were well-worn by late-viii, at which time many fresh butterflies appeared. It is with 
this late emergence that many very dark brownish red individuals appeared at Great 
Swamp. Worn individuals flew until late-ix. A large number of aberrant forms were 
sighted throughout the state in 1983. Of the approximately 50 sightings of archippus 
recorded throughout the year, about 34 can be considered within the normal range of 
variation, with the remainder all of some aberrant nature. Form lanthanis was the most 
common, with 8 sighted or captured. Lanthanis varied considerably as well, from very 
light orange to darker than average ground color. A few specimens have remnant traces 
of the hindwing band. Five of the dark brownish red variety were observed at Great 
Swamp. Two specimens were taken which resemble individuals of the southern subspe- 
cies watsonii, with dark brownish red forewings and pale orange hindwings. Also, one 
dwarf was sighted in West Warwick, but with normal coloration. Areas 1, 2, 3, 4, 6, 8, 
11, 12. (PKW).* 

Limenitis arthemis astyanax Fabricius: Generally uncommon, from mid-vi through 
viii. However, many were observed at Arcadia Wildlife Management Area in mid-viii, 
where one individual of form proserpina was sighted. Another proserpina, with the 
hindwing band greenish, and with a trace white band on the upper forewings, was found 
in central Coventry. One giant female was spotted, perhaps as large as a swallowtail, for 
which it was first mistaken. Areas 1, 2, 3, 4, 8, 12. (KW). 

Nymphalis antiopa Linnaeus: Worn hibernators of the previous fall brood emerged 
in mass-numbers at Great Swamp in early-iv. These were dark, with the marginal bands 
appearing whitish and narrow. Numbers decreased sharply by 1-v, with stragglers on 
the wing until vi. A very small summer brood of 6 was observed elsewhere in the state 
throughout vii. These were very brightly colored, with wide ochreous marginal bands. 
No sightings of the fall brood were made anywhere in the state in 1984. Areas 1, 2, 4, 
8, 11. (KW).* 

Phyciodes tharos tharos Drury: First brood in large numbers throughout vi. Most were 
typical summer form morpheus. While no distinct marcia individuals were identified 
out of hundreds, many morpheus/marcia intermediates were evident. Morpheus was 
abundant in the summer brood from mid-vii through viii. A partial third brood of small 
numbers appeared in mid-ix. These were all morpheus/marcia intermediates much like 


24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


spring specimens. This species was found generally everywhere in weed fields. Areas 1, 
2, 3, 4, 6, 8. (PKW).* 

Polygonia comma Harris: Only three sightings of form harrissi in widely separated 
locations in late-viii and early-ix. Found in woodlands near water. Areas 3, 4, 12. (W).* 

Polygonia interrogationis Fabricius: Surprisingly, only two sightings were made, both 
of form umbrosa. Found along railroad tracks in West Warwick. Late-vi and early-vii. 
Area 2. (K).* 

Junonia coenia Hiibner: A latecomer in late-ix and early-x. Found in small numbers 
only along the coast. None were of any distinct form, but rather, were intermediate 
between wet and dry forms. Areas 9, 10. (WN).* 

Speyeria cybele cybele Fabricius: Surprisingly few sightings, and only at Great Swamp. 
Late-vii. Area 4. (W). 

Vanessa atalanta Linnaeus: Few sightings, most occurring along the coast, but one 
sighting inland at West Warwick. Two broods, first in vii, again in late-ix. Areas 2, 5, 8, 
9. (KNW).* 

Vanessa cardui Linnaeus: Northward migrants and/or local hibernators in late-iv, 
lingering into vi. Summer brood was abundant at inland locations in vii, disappearing 
by viii. Small numbers reappeared briefly in late-viii, with the final brood appearing in 
late-ix to early-x, mostly along the coast. Another common weed-field species. Areas 2, 
4, 5, 8, 9, 10. (KNW).* 

Vanessa virginiensis Drury: An erratic brood sequence was recorded. Small numbers 
appeared in scattered locations during the first week in v, and again during the second 
week in vi. Found throughout vii in small numbers but common only during the third 
week. Small numbers reappeared briefly in late-viii. The final brood appeared during 
late-ix, when this species became fairly common. Generally found in weed fields. Areas 
L, 2,3, 4, 5, 8, 9, 12, (KNW))* 

Papilio glaucus glaucus Linnaeus: Spring “form” canadensis was found in very small 
numbers in the center of the state, in late-iv and early-v. The summer form was recorded 
as occasional sightings in widely scattered locations, throughout vi to viii. I suspect 
canadensis is possibly a univoltine sibling species, but more research is needed. Cana- 
densis in areas 1, 2. (K).* Summer form in areas 1, 2, 4, 5, 8, 11, 12. (KNW).* 

Papilio polyxenes asterius Stoll: Only two spring brood sightings in early-vi. Summer 
brood appeared in small numbers at Great Swamp in late-viii. Areas 2, 4, 5. (KW).* 

Papilio troilus troilus Linnaeus: Spring brood indicated by only one observation in 
early-vi. Summer form sighted at widely scattered locations from mid-vi through viii, 
but only common at Great Swamp in early-viii. Areas 1, 2, 3, 4, 12. (KW).* 

Colias eurytheme Boisduval: Spring form ariadne appeared in small numbers through- 
out v. Summer form amphidusa appeared in small numbers in early-vi. Amphidusa was 
widespread, and common in many areas, throughout vii, but very few were evident in 
viii. The final brood of amphidusa was widespread also from early-ix through x. Several 
different forms of possible eurytheme/philodice hybrids were captured throughout the 
season. One aberrant yellow female was taken at East Matunuck Beach in mid-vi. This 
species can almost always be expected in weed fields. Areas 2, 3, 4, 5, 7, 8, 9, 10, 11, 12. 
(BKNPW).* 

Colias philodice Latreille: Possibly the most persistently abundant species in many 
areas of the state, with three distinct broods of the summer form. Spring form anthyale 
was evident in small numbers only at the onset of the first brood. Common throughout 
v and early-vi, again throughout vii, with an extended final brood from early-viii through 
x. Areas 1, 2, 3, 4, 5, 7, 8, 9, 10, 11, 12. (BKNPW).* 

Pieris rapae Linnaeus: The most widespread species in the state, common in almost 
all open areas but never as abundant as Colias philodice in any area. Spring form metra 
in late-iv. The typical form appeared in 4 broods from vi through x, with the final brood 
being the most common. Areas 1, 2, 3, 4, 5, 7, 9, 10, 11, 12. (BKNPW).* 

Cercyonis pegala alope Fabricius: Very few of these were taken in two locations at 
the western edge of the state in early-viii, with main colonies not being found. The 
distinctive forewing patch is yellow in this subspecies. Areas 3, 6. (PW).* 

Cercyonis pegala maritima Edwards: Abundant in widely scattered colonies from mid- 


VOLUME 39, NUMBER 1 25 


vii through viii. Never before have I observed any butterfly in such large numbers as 
this, at Trustom Pond in South Kingstown, where approximately over one hundred 
individuals were observed along a 30-meter length of trail through a scrub oak thicket. 
The distinctive forewing patch is orange in this subspecies. Maritima also differs from 
subspecies alope by its slightly darker ground color. Differences between these two sub- 
species can only be safely concluded through large series of specimens. Found in grassy/ 
shrubby areas. Areas 1, 2, 4, 8. (KW).* 

Megisto cymela cymela Cramer: Usually found in small numbers in scattered locations, 
preferring grassy open woods. The first appearance was in early-vi, only at Great Swamp. 
Absent in late-vi but reappearing elsewhere in moderate numbers during the first week 
in vii and then in small numbers until early-viii. Areas 1, 2, 3, 4, 5, 8. (KW).* 

Satyrodes appalachia Chermock: Two sightings only at Great Swamp in mid-vii. Sight- 
ed in woods. Area 4. (W).* 


Journal of the Lepidopterists’ Society 
39(1), 1985, 26-32 


FOREST TORTRICIDS TRAPPED USING EUCOSMA AND 
RHYACIONIA SYNTHETIC SEX ATTRACTANTS 


R. E. STEVENS,’ C. SARTWELL,”? T. W. KOERBER,’ J. A. POWELL,?* 
G. E. DATERMAN,? AND L. L. SOWER? 


ABSTRACT. Moths of 31 non-target species of Tortricidae (30 Olethreutinae, 1 Tor- 
tricinae) were lured to synthetic Eucosma and Rhyacionia sex attractants deployed in 
pine forests throughout 12 states in the western U.S. Genera represented include Petrova, 
Barbara, Phaneta, Eucosma, Epiblema, Epinotia, Ancylis, Dichrorampha, Sereda, 
Grapholita, Cydia, and Decodes, as well as a new genus near Rhyacionia. 


In 1977 and 1978 we conducted an extensive trapping survey in pine 
forests in the western United States, using synthetic sex attractants. The 
primary objective was to learn more about geographical distribution 
and host relationships of Eucosma sonomana Kearfott and species of 
Rhyacionia. While the major results have been published (Sartwell et 
al., 1980; Stevens et al., 1980), a variety of other moths, largely ole- 
threutines, also responded to the lures. These were saved and identified 
when their numbers indicated more than chance captures. Generally, 
a minimum of 4-6 similar moths at a trapping location was considered 
sufficient to indicate attraction was not incidental, although in some 
instances we recovered fewer. The catches reported here provide clues 
regarding pheromone chemistry of and possible taxonomic relation- 
ships among certain species. The information may be useful for future 
studies on these and related species. It also provides range extensions 
for some of the species captured. 


METHODS 


Details of the methods, including trapping periods and precise trap 
locations for most collections, are presented in the previously cited 
works (Sartwell et al., 1980; Stevens et al., 1980). In cases in which 
these are not cited, we provide precise trap locations in Table 1. Trap- 
ping periods were similar to those at nearby localities. Briefly, we 
deployed attractant-baited traps in pine forests in most of the western 
United States in spring and early summer 1977 and 1978. Four baits 
were used: (E)-9 dodecenyl acetate (referred to hereafter as (E-9)); 
(Z)-9 dodecenyl acetate (Z-9); a 1:1 mixture of E-9 and Z-9 (50-50); 
and (E,E)-8,10 dodecadieny] acetate (E,E-8,10). 


‘Rocky Mountain Forest and Range Experiment Station, USDA Forest Service, Fort Collins, Colorado 80526. Head- 
quarters at Fort Collins in cooperation with Colorado State University. Present address: Department of Entomology, 
Colorado State University, Fort Collins, Colorado 80523. 

? Pacific Northwest Forest and Range Experiment Station, USDA Forest Service, Corvallis, Oregon 97331. 

° Pacific Southwest Forest and Range Experiment Station, P. O. Box 245, Berkeley, California 94701. 

* Division of Entomology and Parasitology, University of California, Berkeley, California 94720. 


VOLUME 39, NUMBER 1 OT 


The baits were formulated in 3 X 5 mm cylindrical polyvinyl chlo- 
ride pellets containing 4% attractant by weight (Daterman, 1974). Baits 
were mounted on insect pins inserted centrally within Pherocon-II® 
traps. Traps were hung on tree limbs 1.5-2 m above ground. 

Traps were set out at each location in 8 clusters of 5 traps each (4 
baited, 1 unbaited), with no trap nearer than ca. 20 m to another. 
Thus, at each location, each attractant material was presented in 3 
traps, along with 3 unbaited traps used as checks. Trapping periods 
ranged from overnight to more than a month (Stevens et al., 1980). In 
general, traps were deployed 2 to 8 weeks. 

Recovered traps were stored in freezers. Moths were separated by 
presumed species. Representative individuals were removed with for- 
ceps, rinsed in xylene and then hexane or ether to remove the trapping 
adhesive, and finally relaxed and prepared for identification. In some 
instances small parts of traps holding specimens or entire traps were 
immersed in solvent to free specimens. 

Representative specimens are kept in the insect museum at Colorado 
State University, Fort Collins, and at the Essig Museum of Entomology, 
University of California, Berkeley. 


RESULTS AND DISCUSSION 


The species trapped, lures responded to, and localities are presented 
in Table 1. The equivocal nature of a few of the species determinations 
reflects the difficult taxonomic situation in some Olethreutinae. The 
unbaited traps captured only occasional stray moths and were not con- 
sidered attractive. 

Altogether, 31 taxa, all but one being Olethreutinae, are represented 
in the trapped material. Comparing our material with information 
summarized by Inscoe (1982) and Roelofs and Brown (1982), five gen- 
era, Decodes, Ancylis, Phaneta, and Sereda, and a new eucosmine 
near Rhyacionia, are not reported to have been previously captured 
using synthetic attractants. 

Most of the trapped Eucosmini responded to Z-9, E-9, and the 1:1 
mixture of the two (50:50). Phaneta columbiana (Walsingham) was an 
exception, responding almost exclusively to E,E-8,10. Moths of the new 
genus near Rhyacionia responded to E-9, Z-9, and 50-50 except at 
Kingman, Ariz., where all 17 specimens were lured to E,E-8,10. This 
anomaly seems to justify further attention. 

In general, members of the Grapholitini were attracted to E,E-8,10. 
The species of Cydia showed some variability in their responses to the 
preferred materials, reflecting the difficulties in their perceived taxo- 
nomic relationships. C. tana (Kft.) responded to 50-50, but since a total 
of only four specimens were collected this may not be meaningful. 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


However moths of the C. piperana complex were lured in large num- 
bers to 50-50, as well as to E-9 and Z-9 (none to E,E-8,10); this may 
imply some divergence between this complex and other elements of 
the genus. 

The two species of Grapholita we collected responded solely to E,E- 
8,10; more G. conversana Walsingham moths were trapped than any 
other species aside from the target Rhyacionia. By contrast, the much- 
studied oriental fruit moth, G. molesta (Busck) appears to respond 
primarily to compounds unsaturated only at the 8th position in the 12- 
carbon chain (Inscoe, 1982). 


ACKNOWLEDGMENTS 


We thank Richard C. Brown for determining Epinotia species, and J. W. Brewer and 
W. E. Miller for helpful suggestions on the manuscript. 


LITERATURE CITED 


DATERMAN, G. E. 1974. Synthetic sex pheromone for detection survey of European 
pine shoot moth. Res. Pap. U.S. Dep. Agric. For. Serv. PNW-180. 12 pp. 

INSCOE, May N. 1982. Insect attractants, attractant pheromones, and related com- 
pounds. Pp. 201-295, in A. F. Kydonieus & Morton Beroza (eds.). Insect suppression 
with controlled release pheromone systems. CRC Press, Boca Raton, Florida. 

ROELOFS, WENDELL L. & RICHARD L. BROWN. 1982. Pheromones and evolutionary 
relationships of Tortricidae. Ann. Rev. Ecol. Syst. 13:395-422. 

SARTWELL, CHARLES, G. E. DATERMAN, T. W. KOERBER, R. E. STEVENS & L. L. SOWER. 
1980. Distribution and hosts of Eucosma sonomana in the western United States as 
determined by trapping with synthetic sex attractants. Ann. Entomol. Soc. Amer. 
73:254-256. 

STEVENS, ROBERT E., CHARLES SARTWELL, THOMAS W. KOERBER, GARY E. DATERMAN, 
LONNE L. SOWER & JERRY A. POWELL. 1980. Western Rhyacionia (Lepidoptera: 
Tortricidae, Olethreutinae) pine tip moths trapped using synthetic sex attractants. 
Can. Entomol. 112:591-603. 


VOLUME 39, NUMBER 1 


TABLE l. 


29 


Tortricidae other than Eucosma sonomana and Rhyacionia spp. trapped 


with synthetic sex attractants, western United States, 1977 and 1978. 


Species 


Olethreutinae 


Eucosmini—new genus 
near Rhyacionia 


Petrova metallica (Busck) 


Petrova picicolana (Dyar) 


Barbara colfaxiana (Kear- 
fott) 


Phaneta columbiana 
(Walsingham) 


Eucosma bobana Kearfott 


Eucosma ponderosa 
Powell 


Attractant(s) and 


number moths trapped 


E-9 (338) 

Z-9 (3) 

50-50 (32) 
E,E-8,10 (17)? 
E-9 (96) 

Z-9 (16) 
50-50 (45) 
E,E-8,10 (6) 


E-9 (27) 
Z-9 (5) 
50-50 (255) 
E,E-8,10 (1) 


Z-9 (24) 


Z-9 (1) 
E,E-8,10 (43) 


E-9 (1) 

Z-9 (343) 
50-50 (168) 
E,E-8,10 (5) 


Z-9 (14) 
50-50 (8) 


Localities collected! 


Arizona: Kingman A, Portal 
New Mexico: Reserve, Ruidoso, 
Santa Fe, Silver City 


California: Scott Valley (4 km W), 
Tuolumne Mdws., Yosemite Vil- 
lage (12 km NW) 

Nebraska: Alliance 

Oregon: Burns, Crescent Lake, 
Grants Pass, Keno, Sisters 

Washington: Entiat 

Wyoming: Kemmerer 


California: Old Station (10 km 
SE), Sierra City (12 km N), 
Truckee A, Tuolumne Mdws., 
Yosemite Village (12 km NW) 

Colorado: Fraser 

Idaho: Coeur d Alene, Headquar- 
ters 

Montana: East Glacier 

Oregon: Baker, Crater Lake, Cres- 
cent Lake, Idanha, Tiller, Sis- 
ters, Ashland 

Utah: Vernal 

Washington: Leavenworth 

Wyoming: Afton 

Idaho: Headquarters, Coeur 
d'Alene 

Nevada: Las Vegas A 

Oregon: Idanha 

Washington: Kettle Falls 

Wyoming: Afton 

Idaho: Arco 

Nevada: Austin 

Oregon: Lakeview 

Utah: Dutch John 


Arizona: Kingman B 

California: Big Bear City B, Lee 
Vining (4 km SW) 

Colorado: Ft. Collins C, Woodland 
Park 

Idaho: Arco 

Montana: Wolf Creek 

Nevada: Austin, Las Vegas A 

Utah: Dutch John 

Wyoming: Kemmerer 

Oregon: Bly, Chiloquin, Bend, 
Lakeview, Gold Beach, Sisters, 


TABLE l. 


Continued. 


Species 


Eucosma recissoriana 
complex 


Epiblema resumptana 
Walker 

Epinotia emarginana 
(Walsingham) 

Epinotia miscana (Kear- 
fott) 


Epinotia columbia (Kear- 
fott) 


Epinotia n. sp. 
Ancylis columbiana Mc- 
Dunnough 


Ancylis simuloides Mc- 
Dunnough 

Ancylis albafascia Hein- 
rich 

Ancylis pacificana (Wal- 
singham) 

Ancylis mediofasciana 
(Clemens) 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Attractant(s) and 


number moths trapped 


E,E-8,10 (2) 


Z-9 (256) 
50-50 (127) 


50-50 (8) 
E-9 (1)° 


E-9 (3) 
50-50 (1) 

Z-9 (A) 

E-9 (2) 
50-50 (5) 
E,E-8,10 (1) 
E,E-8,10 (32) 
Z-9 (792) 
50-50 (70) 


Z-9 (191) 
Z-9 (7) 
Z-9 (6) 


E-9 (1714) 


Localities collected! 


O’Brien 

Washington: Entiat, Goldendale 

California: Tioga Pass, Tuolumne 
Mdws., Yosemite Village (12 km 
NW) 

Montana: Boulder 

Oregon: Chiloquin, O’Brien 

Utah: Manila 

Wyoming: Buffalo 

Montana: Havre, Wolf Creek 


California: Monterey 
Oregon: Crescent Lake 


Idaho: Headquarters 
Montana: East Glacier 


Arizona: Kingman B 


California: Camino, Descanso (14 
km NE), Emigrant Gap, Fall 
River Mills (4 km NW), Julian 
(3 km SW), Lake Arrowhead (3 
km NE), Monterey, Sierra City 
(12 km N), Upper Lake (20 km 
N) 

Oregon: Bly, Ashland, Cave Jct., 
Crater Lake, Crescent Lake, 
Keno, Idanha, Tiller, Oakridge, 
Sisters, Grants Pass, O’Brien 

Washington: Goldendale, Kettle 
Falls, Leavenworth 


California: Lake Arrowhead, Old 
Station (10 km SE) 


Oregon: Crescent Lake 
Oregon: Bend 


California: Big Bear City A, 
Crestline (1 km NE), Descanso 
(14 km NE), Emigrant Gap, 
Fall River Mills (4 km NW), 
Hat Creek, Julian (3 km SW), 
June Lake (6 km E), Lake Ar- 
rowhead (3 km E), Mt. Shasta 
City, Old Station (10 km SE), 
Placerville, Scott Valley (4 km 
NW), Truckee A, Upper Lake 
(20 km N) 


VOLUME 39, NUMBER 1 ol 


TABLE 1. Continued. 


Attractant(s) and 
Species number moths trapped Localities collected! 


Colorado: Ft. Collins 

Idaho: Arco, Boise, Coeur d’Alene 

Montana: Boulder, East Glacier, 
Superior, Wolf Creek 

Oregon: Ashland, Baker, Bookings, 
Burns, Cave Jct., Crater Lake, 
Crescent Lake, Gold Beach, 
Grants Pass, Idanha, Keno, Lake- 
view, Oakridge, O’Brien, Sisters, 
Tiller 

Utah: Dutch John, Manila 

Washington: Goldendale, Kettle 
Falls, Okanogan, Pomeroy, Shel- 
ton 

Wyoming: Afton, Kemmerer 


Laspeyresiini 

Dichrorampha sedatana 50-50 (6) Montana: Wolf Creek 
(Busck) E,E-8,10 (28) South Dakota: Deadwood 

Sereda tautana (Clemens) _E,E-8,10 (89) New Mexico: Ruidoso, Silver City 

Grapholita caeruleana E,E-8,10 (44) Idaho: Arco 
Walsingham 

Grapholita conversana E,E-8,10 Arizona: Williams (20 km S) 
Walsingham (ca. 2900) California: Clear Lake (20 km N), 


Crestline (1 km NE), Descanso 
(14 km NE), Emigrant Gap, 
Hat Creek, Julian (3 km SW), 
Lake Arrowhead (3 km NE), 
Monterey, Old Station, Sierra 
City (12 km N) 

Colorado: Pagosa Springs 

Idaho: Boise, Headquarters 

Montana: Conner, Havre 

Oregon: Baker, Burns, Cave Jct., 
Crater Lake, Keno, Lakeview, 
Tiller 

South Dakota: Deadwood 

Utah: Cedar City, Vernal 

Washington: Entiat, Goldendale, 
Kettle Falls, Leavenworth, Pom- 
eroy, Tacoma 

Wyoming: Afton, Buffalo 


Cydia populana (Busck) E,E-8,10 (25) Colorado: Redfeather Lakes 
Utah: Dutch John, Manila 


Cydia n. sp. nr. strobilella E,E-8,10 (13) Idaho: Boise 


(L.) 
Cydia tana (Kearfott) 50-50 (4) Oregon: Ashland 
Cydia nr. leucobasis E,E-8,10 (10) Oregon: Idanha 
(Busck) 
Cydia americana (Wal- E,E-8,10 (61) California: Julian (3 km SW), 


singham) Lake Arrowhead (3 km NE) 


32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Continued. 


Attractant(s) and 
Species number moths trapped Localities collected! 


Idaho: Boise 

Oregon: Burns, Eugene, Tiller 

Washington: Entiat, Goldendale 
Cydia colorana Kearfott E,E-8,10 (13) California: Lee Vining (4 km SW) 

Colorado: Ft. Collins C 


Cydia cupressana Kear- Z-9 (4) California: Monterey 
fott 
Cydia piperana complex E-9 (62) California: Old Station 
Z-9 (14) Oregon: Sisters 
50-50 (210) Montana: Boulder, Conner, 


Havre, Wolf Creek 
Washington: Entiat, Goldendale, 
Kettle Falls, Pomeroy 
Utah: Manila, Vernal 
Wyoming: Afton, Kemmerer 


Tortricinae 
Cnephasiini 
Decodes stevensi Powell Z-9 (163) Colorado: Fort Collins C 


1 See Stevens et al. (1980) for more detailed trapping locations, and trapping dates. Detailed locations given here only 
jf not included in Stevens et al. (1980). 

2 E,E-8,10 only from Kingman, Ariz. 

3 Probably incidental. E. emarginana is a generally abundant species and should have been trapped in greater numbers 
if moths responded to the lures. 


Journal of the Lepidopterists’ Society 
$9(1), 1985, 33-42 


NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF 
OPSIPHANES QUITERIA QUIRINUS GODMAN AND 
ERYPHANIS AESACUS BUBOCULUS 
BUTLER (BRASSOLIDAE) 


ROLANDO CUBERO 
C.4, Avs 2-4, Barva-Heredia, COSTA RICA 


ABSTRACT. Details of the various life stages are presented for Opsiphanes quiteria 
quirinus Godman and Eryphanis aesacus buboculus Butler. These are two of the rarest 
Costa Rican brassolids, and this is probably the first photo-illustrated report on their life 
cycles. 


This is the first article of a proposed series devoted to describing the 
early stages, larval host plants and adult behavior of the Brassolidae 
known to occur in Costa Rica. The object of these studies is to provide 
general information and a taxonomic guide to the eight genera (Bras- 
solis, Caligo, Catoblepia, Dynastor, Eryphanis, Narope, Opoptera and 
Opsiphanes) of Brassolidae containing the 19 species reported from 
this country. 


General Descriptions of Life Stages 
Opsiphanes quiteria quirinus Godman 


Egg (Fig. 1). Spherical, slightly flattened at base with numerous vertical ribs; diameter 
ca. 1.5 mm; light green when first laid, becoming dark green with three reddish brown 
concentrical circles when fertile; hatches in 12 days. 

First instar larva (Fig. 2). Head capsule very dark brown, covered with small dark 
brown hairs and setae. Body cylindrical, cream-white in color with several longitudinal 
red bands and reddish brown forked tail. After feeding, color of body changes to light 
green with dark green longitudinal bands; forked tail turns to dark brown. Ca. 15 mm 
long. Moults in 10 days. 

Second instar larva (Fig. 3). Head dark brown with yellow frontal patch on epicranial 
suture; bears four dark brown prominent projections or “horns” along superior and lateral 
borders; head also with whitish pubescence and pair of forward hair-tufts in blackish 
brown color appearing as conspicuous “mustache.” Body with broad longitudinal lemon- 
yellow band on dorsum in quick succession with dark green and light green lateral stripes. 
Anal fork greenish gray. Ca. 24 mm long. Moults in 10 days. 

Third instar larva (Fig. 4). Head with all four horns more developed, with vestigial 
horn at their bases. Yellow frontal patch now broader than in second instar and hair- 
tufts very conspicuous. Body retains dorsal yellow band, but other stripes turn to yellowish 
green and bluish gray. Ca. 39 mm long. Moults in 13 days. 

Fourth instar larva (Fig. 5). Head capsule retains basically color pattern but now horns 
present light orange color with black tips. Dorsal yellow band of body divided by two 
fine light green stripes and rest of body retains same color of previous instar. Ca. 53 mm 
long. Moults in 18 days. 

Fifth instar larva (Fig. 6). Shape of head, as in preceding three instars, basically 
rectangular, with broad pale yellow band along epicranial suture, in quick succession to 
very broad chestnut brown longitudinal band on each epicranium. Color pattern of body 
similar to previous instar; dorsal band now becoming lemon-yellow; spiracles orange- 
brown, and forked tail yellowish green. Ca. 104 mm long. Duration: 16 days. 


34 JOURNAL, OF THE LEPIDOPTERISTS SOCIETY 








Fics. 1-4. Opsiphanes quiteria quirinus: 1, egg, ca. 1.5 mm diameter; 2, first instar 
larva, ca. 15 mm long; 3, second instar larva, ca. 24 mm long; 4, third instar larva, ca. 
39 mm long. 


Prepupa. Prepupal larva loses color pattern of body and changes to translucent green. 
Ca. 65 mm long. Duration: 2 days. 

Pupa (Fig. 7). Pupa grass green; “boat shaped,” laterally compressed at wing cases, 
with head slightly bifid; thin brownish green lines, one along dorsal angle, another lateral 
from cremaster to middle of wing case and another ventral line from cremaster to end 
of antennae; dorsal line gives rise to number of lateral lines in angle directed posteriorly 
to spiracular border; spiracles pale orange; each wing case ventrally with small dark 
brown spot and laterally with golden spot. Ca. 45 mm long. emergence in 22 days. 

Adult. Male (Figs. 8, 9). Body very robust and brownish red. Forewing blackish brown 
with ochreous-orange transverse band and two white subapical spots; basal area reddish 
brown. No pupilate subapical ocelli and three undulate blackish brown submarginal lines, 
with a complicated design in discocellular area; posterior part of ochre-orange band 
clearly visible near margin between median veins. Hindwing brick-red with three ochre- 
orange spots between radials and before terminal margin, which is dark brown; outer 
margin dark brown and very dentate; under surface darker; large spot on radial sector 
and smaller one, greenish ochre, near anal angle; entire surface of wing covered by very 
complicated design of black, brown and yellow-ochre lines and patches which turn darker 
near outer margin; discal cell with conspicuous black-brown spot, which is most char- 
acteristic detail of total design. 

Female (Figs. 10, 11). Forewing blackish brown with white subapical spots larger than 
in male; outer margin somewhat undulated; transverse band very wide and of glossy 
cream-white color; basal area with reddish brown color and hindwing, which is more 
rounded in shape, with same color as male; under surface somewhat paler than male; 
band of forewing entirely visible; but hindwing paler than on male, yet with basically 
same pattern, except in black-brown patch of discal cell, which is more reduced. Span: 
Ca. 80 mm, male and 97 mm, female. 


Eryphanis aesacus buboculus Butler 


Egg (Fig. 12). Round and white when first laid, with numerous ribs; when fertile, turns 
to rosy white color with broad maroon ring, which is not closed at one side for space of 
% mm. Diameter ca. 2.3. mm. Hatches in 12 days. 


VOLUME 39, NUMBER 1 35 





Fics. 5-7. Opsiphanes quiteria quirinus: 5, fourth instar larva, ca. 53 mm long; 6, 
fifth instar larva, ca. 104 mm long; 7, pupa, lateral aspect, ca. 45 mm long. 


First instar larva (Fig. 13). Head rounded, basically reddish brown and covered with 
fine pilosity. Body yellowish orange with red longitudinal bands; with dark brown forked 
tail; ventral surface red. After feeding, bands of body turn very multicolored; with green 
dorsal band limited by two yellow-orange stripes at sides and deep maroon band in quick 
succession with bluish white band on each side. Ca. 18.5 mm long. Moults in 9 days. 

Second instar larva (Fig. 14). Head basically quadrate with three pairs of short in- 
curved horns and particular color pattern of several whitish and greenish brown longi- 
tudinal stripes. Body retains essentially same color and forked tail appears more devel- 
oped. Ca. 28 mm long. Moults in 11 days. 

Third instar larva (Fig. 15). Head with horns more developed showing finely textured 
surface; whitish and greenish brown color pattern persists. Body retains same color as in 
second instar but now there appears four or five fleshy points along dorsum. Ca. 44 mm 
long. Moults in 9 days. 

Fourth instar larva (Fig. 16). Head retains same shape and color pattern. Body brown- 
ish white with broad dorsal line of greenish brown; retaining false spines on dorsum, but 
now more developed and skin (including forked tail), with very particular texture of 
tiny protuberances resembling “chicken skin”; between dorsal line and spiracular bands, 
body also with pair of thin longitudinal dark brown stripes. Ca. 80 mm long. Moults in 
11 days. 

Fifth instar larva (Fig. 17). Head large and broad; dirty white in color but now covered 
with fine pilosity. Body light brown-yellow, slender striped with greenish brown lines 
and longitudinal row of short dark brown lines on each side, just beyond spiracular band; 
below this band, larva with longitudinal border of short hair, lighter in color than on rest 
of body; ventral surface reddish brown. During this instar red scent gland of larva, located 
in front of the prothoraxic legs, particularly visible. Ca. 118 mm long. Duration 26 days. 

Prepupa. Prepupal stage very short, requiring two days; characterized by paler and 
semi-translucent color of body which becomes shorter than fifth instar, larva adopts 
hanging position for pupation usually attached at tip of fresh rolled leaf of host plant. 

Pupa (Fig. 18). Elongated, dirty rosy white with fine pattern of pale grey-brown stripes 
and patches resembling dried and rolled leaf of bamboo; abdominal segments thickening 
gradually from Al10 to A4; wing cases very compressed, extending from Tl to A4; 
dorsally, pupa with diffuse grey-brown line and row of fine dark brown points at each 


36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


side; these points particularly visible on Al to A4; also, with two more longitudinal lines 
of same color, one subdorsally with dark brown spot on A4 and another laterally just 
along spiracular band; cremaster and head cristulae appear darker than rest of pupa; 
head cristulae apposed and somewhat incurved ventrally; middle of each wing case and 
just over lateral stripe with small and somewhat diffuse yellow-ochre spot. Ca. 56 mm 
long. Adult emerges in 20 days. 

Adult. Male (Figs. 19, 20). Forewing blackish brown with median area of blue-violet 
tone; tone also present on hindwing forming band along submarginal area, from subcostal 
vein to middle of wing and small patch on discal section. Hindwing also with rounded 
yellowish spot at inner margin with scent scales and black patch between this spot and 
end of blue-violet submarginal band. 

Female (Fig. 21). Paler than male. Forewing with ochreous submarginal band forked 
anteriorly; with very translucent zone from basal to submedian area and design of un- 
derside clearly visible; this zone with pale blue-grey reflection; this pigmentation occur- 
ring from the basal to the median area, delimited by broad bluish-brown band from 
postmedian to outer margin; the inner margin with pale brownish gray color; on under- 
side (Fig. 22), color pattern between male and female very similar, basically with details 
that permit identification of this species from others of genus, such as well marked and 
colored designs of gray, black, white and ochre patches and lines and presence of double 
eyespot on hindwing. Wingspan: 170 mm, male; 112 mm, female. 


Natural History 


According to Fruhstorfer (in Seitz, 1924), Opsiphanes quiteria qui- 
rinus Godman occurs from Guatemala to Panama; nevertheless, it is 
considered a very rare butterfly. In Costa Rica this species occurs from 
sea level to about 1000 m in the wet forests but is very seldom seen. 
It flies generally in the afternoon, mostly in the forests, but sometimes 
it is possible to observe a specimen flying across open areas. 

O. quiteria quirinus represents one of the largest species of the Costa 
Rican Opsiphanes, occurring together with O. cassina fabricii Bois- 
duval and O. invirae cuspidatus Stichel, and they probably share larval 
food plants (those in the Palmae family). I once saw a female of the 
former species ovipositing on a palm of Geonoma sp. in an inaccessible 
place in the forest at Estacion Agua Fria at the Tortuguero National 
Park in the Province of Lim6én (20 m) and again at Colonia Virgen del 
Socorro, Province of Heredia (800 m), on the palm, Prestoea allenii 
H. E. More. According to these observations the female lays the eggs 
singly on the leaf but about four to six per plant. The larva is very 
cryptic while on the host plant, resting on the underside of the leaf 
during the first and second instars and later, in a “silk bed” that the 
caterpillar constructs with the semi-rolled tip of the leaf. 

The adult is a very fast flier and feeds (as do all the species of the 
genus) on sap and rotting or decaying fruits. 

Eryphanis aesacus buboculus Butler has a wide distribution in the 
Costa Rican forests, and it occurs from 500 to about 1200 m. Never- 
theless, with this wide distribution it represents a rare species with very 
restricted flying areas in association with water courses and bamboo 
thickets. Little is known of the adult behavior of this butterfly, but 


VOLUME 39, NUMBER 1 37 





Figed Fige9 





Fige11 


Fics. 8-11. Opsiphanes quiteria quirinus: 8 & 9, male, dorsal and ventral aspects, 
80 mm wingspan; 10 & 11, female, dorsal and ventral aspects, 95 mm wingspan. 


some observations, principally at Hacienda Santa Maria at the Rincon 
de la Vieja National Park in the Province of Guanacaste (900 m), 
reveals that E. aesacus buboculus is basically a crepuscular flier. It flies 
at dusk, essentially a courtship activity, which is characterized by ter- 
ritoriality behavior of the males along the river edges. The blue-violet 
iridescence of the male wings during their rapid flight is very conspic- 
uous. The female receives the courting male on the surface of broad 
leaves, always on the river edges and particularly near the water falls. 
They are active at dusk for a period of about one hour and curiously 
halt all activity and disappear when bats begin to fly. 

I suspect, as a result of a number of experiences using butterfly traps 
and banana bait, that the feeding habits take place mainly in the 
morning. E. aesacus buboculus feeds on sap and rotting fruits. I ob- 
served specimens feeding on decaying fruits, and only once did I see 
a female feeding on a bunch of fruits of Ruagea caoba (C. DC.) Harms 
(Meliaceae), approximately four meters above the ground near the Rio 
Sarapiqui bridge on the way to Colonia Virgen del Socorro. 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





a 


Figet3 v aa 





Fics. 12-15. Eryphanis aesacus buboculus: 12, eggs on Bambusa vulgaris; 13, first 
instar larva after feeding, ca. 18.5 mm long; 14, second instar larva, ca. 28 mm long; 
15, third instar larva in resting position, ca. 44 mm long. 


My observations of the oviposition habits of E. aesacus buboculus 
are restricted exclusively to my greenhouse work. The female used for 
this study laid fourteen eggs, from four to six per day. Eggs were laid 
in pairs or triplets on the leaves of Bambusa vulgaris Schard. ex Wendl. 
(Gramineae), used as a substitute for Olyra caudata Trin. (Gramineae), 
which is the primary natural food plant, at least for the area of Colonia 
Virgen del Socorro; although, it is possible that E. aesacus buboculus 
is using a wide spectrum of host plants of the Gramineae, including 
Chusquea scabra Sods. & Cald., which is also present there. 

The larvae of E. aesacus buboculus are very cryptic when on the 
host plant, especially in the fourth and fifth instars. They rest at the 
end of the leaves and become very agitated when disturbed, separating 
and erecting their forked tail and secreting an odorous substance by 
means of the scent gland located in front of the prothoracic legs. The 
pupa is also very cryptic. Presently, I do not have any information 
about specific or generalized parasites for this brassolid. 


Discussion 


What impressed me is the scarcity of data published about the Bras- 
solidae, especially in respect to the description of early stages and food 
plant records. It represents a serious problem for the correct identifi- 


VOLUME 39, NUMBER 1 39 





Fics. 16-18. Eryphanis aesacus buboculus: 16, fourth instar larva, ca. 80 mm long; 
17, fifth instar larva, ca. 118 mm long; 18, pupa, lateral aspect, ca. 56 mm long. 


cation of the species in a given locality. I still have many problems 
establishing the exact location of some species in this family, and the 
possibility of obtaining new data on Brassolidae for Central America 
looks almost impossible. Fruhstorfer’s (in Seitz, 1924) data on the early 
stages are still the most complete information available, but they are 
insufficient and some of the adult species descriptions are not precise. 
That is the case with the description of the O. quiteria quirinus female, 
which is different from the original of Godman and Salvin (1879- 
1901). There are some short papers available that offer some help in 
establishing a relationship between Central and South American species, 
with short descriptions of the life cycles as in Rothschild (1916), but it 
is extremely urgent that there be a complete revision of the Brassolidae. 

O. quiteria quirinus feeds on palms, but it is necessary to have more 
data about the species of palms used. I can supply this information as 
regards the palms of the genus Geonoma and Prestoea, in addition to 
a life cycle study on O. invirae cuspidatus using the same host plants. 
Also, there are more complete works on the common O. cassina fabricii 
as in Young and Muyshondt (1975), in which the palms Cocos and 
Bactris are reported as hosts for this species. Therefore, I suggest the 
possibility that O. quiteria quirinus uses a broad spectrum of species 
in the Palmae as host plants. I reared separately three larvae of O. 
quiteria quirinus on Chrisalidocarpus lutescens H. Wendl, which is 
an introduced species of palm (L. J. Poveda, pers. comm.; Holridge 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fig.19 Fige20 


/ , 
VM ‘ 
ae 


. . 
et Phy 





Fig .22 


Fics. 19-22. Eryphanis aesacus buboculus. 19 & 20, male, dorsal and ventral as- 
pects, 107 mm wingspan; 21 & 22, female, dorsal and ventral aspects, 112 mm wingspan. 


and Poveda, 1975), and I obtained the same satisfactory results as on 
Prestoea. I also have C. lutescens at my home in Barva, Province of 
Heredia (1200 m), and it is constantly frequented by O. cassina fe- 
males for oviposition. 

In respect to the Opsiphanes caterpillars, it is necessary to mention 
the paper of Young and Muyshondt (1975), since these authors are in 
error when they predict that the eversible gland located in front of the 
prothoracic legs of Caligo and Morpho caterpillars is absent in Opsi- 
phanes spp. I have observed that this gland is present in O. cassina 
fabricii, O. invirae cuspidatus and O. quiteria quirinus larvae, and is 
probably present in all the Opsiphanes larvae, including those of the 
old description of Miiller (1886), which was disputed in Young and 
Muyshondt’s article. 

Eryphanis aesacus buboculus, according to Fruhstorfer’s report (in 
Seitz, 1924), occurs from Nicaragua to Colombia. The distribution re- 
ports for the Costa Rican area come from very different habitats in 
the North, the Pacific and the Atlantic slopes and to some extent due 
to this circumstance, I suspect that this species uses many genera in 


VOLUME 39, NUMBER 1 4] 


the Gramineae as host plants. According to Poveda (pers. comm.) and 
to Standley’s (1937) report, Olyra caudata is considered a very rare 
species, and the identification of this plant was very difficult. To sup- 
port my theory, I reared some larvae of E. aesacus buboculus on Chus- 
quea scabra with satisfactory results. Also, I received a report from 
Miguel Serrano (pers. comm.) of E. aesacus aesacus H.-Schaffer being 
reared on Bambusa vulgaris in El Salvador during 1967. 

At Colonia Virgen del Socorro, I discovered that E. aesacus bubo- 
culus uses Olyra caudata, together with the satyrid Oxeoschistus puer- 
ta submaculata Butler and Druce, and curiously, this satyrid shares 
Chusquea longifolia Swallen with the brassolid Opoptera staundingeri 
Godman and Salvin at the Monte de la Cruz, Province of Heredia 
(2000 m). I hope that future studies with these species can further help 
us understand their relationships. 

Another case of affinity occurs with the early stages and adult be- 
havior observed for E. aesacus buboculus and E. polyxena lycomedon 
Felder. The only places where I have reports of the co-existence of 
these two species are from P. J. deVries (pers. comm.) at Turrialba, 
Province of Cartago (600 m), and from R. L. Hesterberg (pers. comm.) 
at Finca El Rodeo, Province of San José (500-600 m). Malcolm Barcant 
(1970) gives some short references on the adult behavior of E. polyxena 
polyxena Meerb. from Trinidad-Tobago, and it is also very interesting 
to observe that a close relationship exists between the habits of this 
species and those of the Costa Rican species of Eryphanis. 


ACKNOWLEDGMENTS 


I am very grateful for the kind assistance and support of the following persons and 
institutions: the Servicio de Parques Nacionales (National Park Service) and the Direcci6n 
General Forestal (Forestry Dept.), for their kind permission to work at the study sites 
mentioned in this paper; to Phillip J. DeVries and Isidro Chacén, who gave me the 
opportunity to review the butterfly collection at the Museo Nacional de Costa Rica 
(National Museum); to biologists Luis J. Poveda, Luis Diego Gdmez, and Rafael Ocampo, 
who helped me with the identification of host plants. My special gratitude to Miguel 
Serrano, Richard L. Hesterberg, Rubén Canet, Eugenio Corea and Jim Scionka, who 
offered support with data, field studies, and revision of the present document, and to Dr. 
Lee Miller, Curator of the Allyn Museum of Entomology (Florida) for assistance with 
the identification of O. quiteria quirinus. 


LITERATURE CITED 


BARCANT, M. 1970. Butterflies of Trinidad and Tobago. Collins-Type Press, London. 
Pp. 117-118. 

GopMAN, F. D. & O. SALVIN. 1879-1901. Biologia Centrali-Americana, Insecta, Lep- 
idoptera—Rhopalocera. Vol. I. 128 pp. 

HOLDRIDGE, L. R. & L. J. POVEDA. 1975. Arboles de Costa Rica, Vol. I. Centro Cien- 
tifico Tropical, San José. 51 pp. 

MULLER, W. 1886. Siidamerikanische Nymphalidenraupen. Zed. Jahrb. Db. 1:417-678. 


42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ROTHSCHILD, W. 1916. Notes on Amathusiidae, Brassolidae, Morphidae, etc. with de- 
scriptions of new forms. Novitates Zoologicae 23:299-318. 

SEITZ, A. 1924. Macrolepidoptera of the world. Vol. 5. The American Rhopalocera. 
Alfred, Kernan Verlag, Stuttgart. Pp. 285-332. 

STANDLEY, P. C. 1937. Flora of Costa Rica. Vol. 17, Part I. Field Museum of Natural 
History, Chicago. 

YOUNG, A. M. & A. MUYSHONDT. 1975. Studies in the natural history in the family 
cluster Satyridae-Brassolidae-Morphidae (Lepidoptera, Nymphaloidea) III. Opsi- 


phanes tamarindi and Opsiphanes cassina in Costa Rica and El Salvador. Studies 
on Neotropical Fauna. Pp. 19-55. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 43-47 


THE DEFENSIVE ENSEMBLES OF TWO PALATABLE MOTHS 


Davip L. EVANS 


Department of Biology, American University of Beirut, 
Beirut, Lebanon 


ABSTRACT. I compared the primary (before disturbance) and secondary resting 
places, relative palatabilities, and escape behaviors of a feces-mimicking moth and a 
specific background matching moth. The components of the defensive ensemble are 
discussed. 


There are two levels of anti-predation mechanisms, primary and 
secondary. Primary defensive strategies are those which are effective 
before the attack of the predator, e.g. background matching, apose- 
matism, dispersion, anachoresis. Secondary techniques include protean 
escapes, noxious discharges, unpalatability, etc. (Edmunds, 1974). Pre- 
viously, I had shown that there are different types of secondary pro- 
tective behaviors which accompany aposematism and crypsis (Evans, 
1983) and crypsis and Batesian mimicry (Evans, 1978). Possibly dif- 
ferent subtypes of crypsis may be accompanied by different types of 
secondary defensive behavior. It is possible to recognize at least two 
types of cryptic organisms. Some animals have a specific color pattern, 
body outline, and behavior which allows them to match a particular 
portion of their habitat (Sargent, 1981). Apparently, there are other 
organisms which because of their generalized dull coloration, can blend 
into several backgrounds but none perfectly. Some authors (Agee, 1969 
with Heliothis zea; Knight, 1916 with Pseudaletia unipuncta) have 
noted moths which rest in more than one situation in their habitats, 
but a similar observation could be made with some large mammals. 

Resemblance to dung might be considered a third type of crypsis. 
Some authorities could argue that dung-like organisms are Batesian 
mimics, since they do not match any background. Cryptics such as 
stone-like plants (e.g. Lithops spp., Mesembryanthaceae) also resemble 
inedible objects but are found among a general background of the type 
of stones they resemble. Conversely, bird dung mimics would always 
be found naturally in ecosystems where bird droppings exist. The two 
basic strategies, i.e. crypsis and Batesian mimicry, seem to be parts of 
a continuum of deception types. 

In this study I wished to compare the primary and secondary pro- 
tective activities of a moth which resembled bird droppings to those 
activities of a moth which seemed to be a good match of a specific 
background. 

Concochares arizonae Hy. Edw. (Noctuidae) is a black and white 
moth which, in its natural resting position, with wings folded, closely 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


resembles a medium-sized passerine (Aves) dropping. It does not match 
the background of its habitat in general but it might seek out those 
specific situations where it would become inapparent. Bomolocha vaga 
(Hiibner) (Noctuidae) is a russet and tan colored moth which seemed 
(to my eyes) successfully cryptic while resting on specific substrates. 
The dorsal coloration of B. vega suggests both the coloration and the 
texture of the broken stones which are abundant in its environment. 


METHODS 


Cryptic animals match their resting backgrounds to the exposed 
color pattern of their bodies (Sargent, 1981), display protean escape 
upon perceiving predator-like stimuli (Evans, 1983; Humphries & 
Driver, 1970), and are palatable (Rothschild, 1981). A study of a cryp- 
tic animal should touch on each of these points. 

I conducted this study on the Yuma (Arizona) Desert near Fortuna 
wash. The area is a mixed Larrea-Cercidium community, the ground 
littered with weathered and broken dark-red stones. I performed the 
experiments from sunrise to one hour later in March when the ambient 
temperature varied 9-17°C and the moisture 18-42% R.H. Both moth 
species were assembled using an ultraviolet light placed on the ground 
in open areas the evening before the tests. I found it necessary to 
change the light’s location three times during the course of the exper- 
iments because the iocal bird community repeatedly began to disturb 
the moths. Similar problems have previously been reported with assem- 
bling palatable moths in natural settings (Jeffords et al., 1979). I tested 
the moths’ reactions to a dorsal front-wing touch using a needle as in 
a previous study (Evans, 1983). The front wing touch mimics the initial 
tentative attack by an avian insectivore. I recorded the reaction du- 
ration (+0.2 s), reaction type, and resting places before and after stim- 
ulus. I captured each moth after testing and maintained it at —20°C 
prior to the palatability tests. 

The palatability of the moths was determined using standard tech- 
niques (Evans & Waldbauer, 1982; Evans, 1984). The test birds were 
three wild-caught adult northern catbirds (Dumetella carolinensis (L.) 
and two brown thrashers (Toxostoma rufum (L.) (both Passeriformes: 
Mimidae). Three acceptances in successive offerings for each bird con- 
stituted evidence of palatability in that species of moth. 

I analyzed the data using r x c contingency tables (Snedecor & 
Cochran, 1980) since there was a non-normal distribution. 


RESULTS 


Both species of birds ate the moths. B. vega was eaten quickly and 
usually in preference to the alternative prey, Tenebrio molitor L. pu- 


VOLUME 39, NUMBER 1 45 


ER 


Omen ort 5h.) 6 | 7 8 1G.) Fo an) I2 49s eas 16. 47) 18). VIG) 20 (21 220 23. 25 


10 


mas) 


{ 
ORewlogec mes tee S56 7) 8. 9101 12. 32 713 44 #15 16 #417 #+218 #49 20 21 22 23 a 25 


Seconds of Flight after Stimlus => 
Fic. 1. Flight duration histogram. Flights timed +0.2 s. Each moth was killed after 
a single test. The upper graph illustrates the flight duration frequencies of Conochares 
arizonae and the bottom graph those of Bomolocha vega (both Lepidoptera: Noctuidae). 
Additional times for C. arizonae not shown: 34.2 and 66.8 s. Additional durations of B. 
vega not shown: 36.8 and 55.8 s. The two moths have statistically significantly different 
flight duration frequencies (P < 0.005, r x c). 


pae (Coleoptera: Tenebrionidae). This moth is doubtless a highly pal- 
atable species: I found numbers of their body-less wings near the light 
later in the day and saw (through a blind) house sparrows (Passer 
domesticus L.) feeding on the insects by first tearing off the moths’ 
wings. The catbirds initially accepted C. arizonae with hesitation and 
after consuming the alternate prey. Upon subsequent presentations, 
the catbirds became more receptive to C. arizonae and began to eat 
them more quickly. Bent (1948) reports that adult female catbirds 
sometimes feed on their nestlings’ droppings. Initially, nestling drop- 
pings are differently shaped than those of adults which C. arizonae 
resembles. The thrashers refused the latter moth species at first, but 
one of the thrashers began to eat them in later presentations. The other 
thrasher never touched the droppings-mimic. All four birds would hold 
this moth in their beaks for some time before ingesting it. 

Near the ultraviolet light, I found all of the B. vega resting initially 
on substrates matching that of the dorsal (exposed) surface of their 
outstretched wings. There was really nothing other than bird feces in 
the area which matched C. arizonae. None of these moths was resting 
on or near bird feces. 

Figure 1 illustrates the flight durations of the two species. I divided 


46 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the 128 flight durations into 7 groups: 0 (no flight), 0.2-0.8 s, 1.0-1.6, 
1.8-2.8, 3.0-4.0, 4.2-5.6, =5.8. The difference between the responses 
in the two moths is statistically significant (P < 0.005). Thirty-nine 
percent of the chi-square value came from the last classification. 

Most of the flying moths changed the direction of flight at least once 
while in the air (B. vega, 84%; C. arizonae, 80%). The brief (S0.4 s) 
flights of the feces mimics, however, were largely (88% of those short 
flights with at least one change in direction) straight away from the 
substrate and almost straight back to a place close to the starting point. 
These moths seemed to land on random objects. B. vega flights were 
typically protean (Roeder & Treat, 1961). When the moths landed, 
82% of the B. vega landed on a matching substrate. The initial location 
of the moth was one that it had made, presumably, without the recent 
stress of the predator-like stimulus. The moths select the secondary 
landing site under potential pressure from a predator so that occasional 
mistakes must be made. I saw a few B. vega change their secondary 
landing site. 


DISCUSSION 


Clearly, the mimic of bird droppings was more likely to make quick- 
er, shorter flights than the specific background matcher, B. vega. It 
seems logical that C. arizonae had brief flights since there was no 
particular background which they matched. In any case, the strategy 
of a fecal mimic is not to blend in exactly with a particular area but 
to be an apparent but ignored object. A bird dropping which flew 
relatively long horizontal distances would attract attention. Conversely, 
the B. vega encountered on their longer flights some non-matching or 
poorly matching substrates (trees, bushes, dead twigs, etc.) which most 
avoided. This activity would be likely to require more time. Appar- 
ently, if the optimal match is not discovered then a poorer match is 
sometimes accepted temporarily. In an earlier study (Evans, 1983), 
another moth resembling an inanimate object, Datana ministra (Dru- 
ry) (dead twig mimic), exhibited a somewhat different pattern of es- 
cape behavior than that of taxonomically related general background 
matchers. Moth flight is ambient temperature dependent; therefore, 
these two studies are not precisely comparable since the present inves- 
tigation was conducted with moths at lower air temperatures. Never- 
theless, it appears that different escape behavior patterns may accom- 
pany different cryptic techniques. There are unified protective 
ensembles, the components of which can be isolated and shown to be 
characteristically distinct from analogous components in other life 
strategies. 


VOLUME 39, NUMBER 1 47 


ACKNOWLEDGMENTS 


I wish to thank my parents, Mr. and Mrs. E. E. Evans for their help while I was in 
Yuma; Drs. Arslanian, Webb, and Waldbauer for their continued encouragement; and 
the School of Arts and Sciences of the American University of Beirut for their financial 


support. 


LITERATURE CITED 


AGEE, H. R. 1969. Mating behavior of bollworm moths. Entomol. Soc. Am. Ann. 62: 
1120-1122. 

BENT, A. C. 1948. Life histories of North American nuthatches, wrens, thrashers, and 
their allies. U.S. Natl. Mus. Bull. 195. 

EDMUNDS, M. 1974. Defence in animals. Longman Group Limited, London. 

Evans, D. L. 1978. Defensive behavior in Callosamia promethea and Hyalophora 
cecropia (Lepidoptera: Saturniidae). Am. Midland Nat. 100:475-479. 

1988. Relative defensive behavior of some moths and the implications to pred- 

ator-prey interactions. Entomol. Exp. et Appl. 33:103-111. 

1984. Reactions of some adult passerines to Bombus pennsylvanicus and its 
mimic, Mallota bautias. Ibis 126:50-58. 

Evans, D. L. & G. P. WALDBAUER. 1982. Behavior of adult and naive birds when 
presented with a bumblebee and its mimic. Z. Tierpsychol. 59:247-259. 

HUMPHRIES, D. A. & P. M. DRIVER. 1970. Protean defense by prey animals. Oecologia 
5:285-302. 

JEFFORDS, M. R., J. G. STERNBURG & G. P. WALDBAUER. 1979. Batesian mimicry: Field 
demonstration of the survival value of pipevine swallow-tail and monarch color 
patterns. Evolution 33:275-286. 

KNIGHT, H. H. 1916. The army-worm in New York in 1914. New York Agr. Exp. Sta. 
Bull. 376:749-765. 

ROEDER, K. D. & A. E. TREAT. 1961. The detection and evasion of bats by moths. Am. 
Sci. 49:135-148. 

ROTHSCHILD, M. 1981. The mimicrats must move with the times. Biological J. Linn. 
Soc. 16:21-23. 

SARGENT, T. D. 1981. Antipredator adaptations of underwing. Pp. 259-284, in A. G. 
Kamil & T. D. Sargent (eds.). Foraging behavior. Garland STPM Press, New York. 

SNEDECOR, G. W. & W. G. COCHRAN. 1980. Statistical methods. 7th ed. The Iowa 
State University Press, Ames, Iowa. 








Journal of the Lepidopterists’ Society 
39(1), 1985, 48-50 


CYMAENES FINCA, SP. N. (HESPERIIDAE) 
FROM TRINIDAD, W.LI. 


M. J. W. Cock 


Commonwealth Institute of Biological Control, Imperial College, 
Silwood Park, Sunninghill, Ascot, Berkshire, U.K. 


ABSTRACT. Cymaenes finca, sp. n. (Lepidoptera: Hesperiidae) is described as new 
from the island of Trinidad, West Indies. The male genitalia and forewing venation and 
markings are illustrated. It is compared with C. tripunctus theogenis Capronnier, and 
the male genitalia of that species are also illustrated. 


Cymaenes finca, sp. n., described below, was taken by the author 
on the island of Trinidad, but not included in the recent list of the 
Hesperiidae of Trinidad and Tobago (Cock, 1982). It is compared with 
C. tripunctus H.-S. and C. lepta Hayward. The abbreviations and 
terminology used follow Evans (1955). 


Cymaenes finca, new species 


Fig. 1 (6 FW); 2-5 (6 genitalia) 


Description. 6 F 15-17 mm. Upf brown; white spot in space 2 below origin of vein 
4; another in space 3, and sometimes a very small one in 6; pale brown spot in space 1B 
faintly apparent and not always present; cilia concolorous (Fig. 1). Uph unmarked, brown; 
darker in space 7 and along termen, cilia concolorous. Unf brown, black on disc; spots 
as for upf; broad pale area in space 1B. Unh pale brown; faint white spots in spaces 2 
and 3; inconspicuous pale streak in space 1C and along vein 2. Antennae dark above; 
barred along front margin of shaft; yellow below base of club; orange-brown below 
apiculus. Palpi paler basally, cheeks pale brown. Thorax and abdomen color match wings, 
except abdomen paler below. Mid tibiae five spines. Genitalia (Figs. 2-5); end of cuiller 
broadened, excavate and doubly spined at tip only. Uncus and gnathos deeply and 
broadly divided viewed dorsally, and deeply divided viewed from side. 

? F 15 mm. In poor condition. Upf with white spots 1B (faint), 2, 3, 6-8; unf pale 
area in 1B inconspicuous; unh pale spots 2-6; otherwise as 6. 

Type material. Holotype male: Trinidad, W.I., Las Lomas, Spanish Farm, 17.XII.1980, 
MJW Cock; allotype female: same locality and collector, 2. VIII.1981; paratype males: 
same locality and collector, 1 6 7.III.1980; 2 44 at dusk, 23.III.1980; 1 6 at dusk, 4.X1.1980; 
1 6 17.XI1.1980; 2 66 2. VIII.1981. 

Deposition of type material. I retain two paratypes, one will be sent to the National 
Museum of Natural History, Washington, and the remaining paratypes, the allotype and 
the holotype will be deposited at the British Museum (Natural History). 


Discussion. The double spined cuiller tip of the male valve puts this 
species in the same group as C. tripunctus and C. lepta. C. finca, 
however, is only doubly spined at the tip of the cuiller. Compared with 
C. tripunctus theogenis Capronnier (Figs. 6-8), C. finca is larger, the 
wings are more produced, and the uncus and gnathos are more strongly 
divided. C. lepta is much more extensively spotted (Evans, 1955), hav- 
ing a “broad continuous row of pale spots from space 1C to 7, broadly 
dark-edged on both sides and a spot before end cell’? on unh and 


VOLUME 39, NUMBER 1 49 





{ iaem | 


Fic. 1. FW venation and UPS markings of 6 Cymaenes finca, sp. n. 


“generally conspicuous white spots in spaces 1B, 2, 3 and 6-8 and may 
be cell dot’ on upf. 

The type locality, Spanish Farm, is a small patch of lowland forest 
on a ridgetop, just north of the road from Las Lomas to San Rafael. 





3 


Fics. 2-5. Male genitalia Cymaenes finca, sp. n. 2, dorsal view uncus and gnathos. 
3, lateral view without claspers. 4, left clasper internal view. 5, left clasper ventral view. 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1 mm 








Fics. 6-8. Male genitalia Cymaenes tripunctus theogenis Capronnier from Trini- 
dad. 6, dorsal view uncus and gnathos. 7, lateral view without claspers. 8, left clasper 
ventral view. 


As Barcant (1970) has pointed out, it harbors a rich butterfly fauna. C. 
finca is the second species to be described from this locality in recent 
years. The riodinid Pachythone barcanti Tite (not Barcant) was also 
described from here, although it is also known from other areas of 
Trinidad, e.g., Sans Souci Estate, Sangre Grande (F. C. Urich, pers. 
comm.). This and the wide variety of interesting and rare species to 
be found there justify efforts to preserve what remains of this small 
wood. 


LITERATURE CITED 


BARCANT, M. 1970. Butterflies of Trinidad and Tobago. Collins, London. 314 pp. 

Cock, M. J. W. 1982. The skipper butterflies (Hesperiidae) of Trinidad. Part II Sys- 
tematic list of the Trinidad and Tobago species. Occas. Papers Dept. Zool., Univ. 
West Indies, St. Augustine, Trinidad, No. 5. 49 pp. 

Evans, W. H. 1955. A catalogue of the American Hesperiidae in the British Museum 
(Natural History). Part IV. Hesperiinae and Megathyminae. British Museum (Nat- 
ural History) Publication. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 51 


BOOK REVIEW 


LEPIDOPTERA: HESPERIIDAE. NOTES ON SPECIES-GROUP NAMES, by Charles A. Bridges. 
Urbana, Illinois, publ. by the author: ii + 1.129 + II.41 + III.62 + IV.30 + V.13 pp. Price: 
$35.00 (available from author: 502 W. Main St., Apt. 120, Urbana, IL 61801). 


This is an impressive compendium of the known literature on the skipper butterflies 
of the world. There is a brief Introduction, 129 pages of Species-group Names of Hes- 
periidae, a 41-page Index of Genera with their included species, an Index of Authors 
and Bibliography (62 pages), a 30-page Bibliography and a 16-page Index to Journals 
and Serials. All of these are very useful to the specialist, and it certainly gives an idea of 
the library that needs to be available for systematic work on a worldwide basis. 

Citations for the names are given in Section I, along with type localities and sites of 
deposition of the type specimens, where these data are known, using a similar format to 
that used by F. M. Brown and myself in our Catalogue /Checklist but much more 
compact. The type localities are abbreviated, however. It is interesting that on page I.1 
the statement is made that the arrangement is based entirely on bibliographic references; 
that no specimens have been examined and that no new names are introduced. Therein, 
to some users, is a problem with this volume: but it is not the province of a bibliographic 
compilation to pass such judgments. One has a right, however, to expect that the biblio- 
graphic citations have been checked for their accuracy. Regrettably, such is not always 
the case in this work. I have closely examined only those citations that impinge on my 
own work and find that there are definite misstatements in the book. 

On page I.2 the citation for Thorybes aemilea (Skinner) is given as an incorrect 
subsequent spelling (ISS) of aemilia, but examination of Skinner’s original description 
shows that the proper spelling is indeed “aemilea.” The aemilia spelling apparently 
dates from Holland’s Butterfly Book in 1898, which was copied by Evans in his Cata- 
logue. There was at least one ISS in the Miller and Brown Catalogue/Checklist that 
Bridges did not catch: Parker described Hesperia powesheik from Grinnell, Iowa in 1870. 
Grinnell is in Poweshiek county, and we spelled the species name the same as the county 
name; I have been unable to find that lapsus elsewhere, and we must be blamed for it. 

Some nomenclatorial errors are also promulgated in this book. For example, on page 
1.60 Hesperia julianus Turton, 1802 is listed under “d” (available, invalid, unused), 
whereas, H. julianus Latreille, [1824] is listed right below it as “c”’ (available, valid 
synonym). Clearly, Latreille created an invalid junior homonym, whereas, Turton’s name 
is indeed available; so, the designations on these two names should be reversed. Elsewhere 
(page 1.15), my subspecies benitoensis is listed as in the genus Unkana, an Indo-Malayan 
genus. My insect was named in the African genus Ceratrichia, and the entire paper was 
devoted to African skippers. The name in question was applied to a subspecies of Cer- 
atrichia flava (Hewitson), rather than Unkana flava Evans from India. C. flava and 
several of its subspecies are mentioned in II.8, and the mixup here is puzzling. 

These complaints are not made to diminish from the book’s great value. They are 
more to alert workers to the necessity of checking original sources in cases of a nomen- 
clatural problem. Secondary sources (of which this book is one) should be used as indi- 
cations of where to find original data and not considered to be the data themselves. In 
this regard, and with the above caveats, I consider this book to be one of the most 
significant weapons in the hesperiid systematist’s arsenal. Bridges has written me that he 
intends a revised edition incorporating the corrections that he receives from other work- 
ers. This work is a great deal better on names than is the average museum card catalogue, 
even in its present form, and workers on groups other than the skippers can only hope 
that Bridges will favor us with volumes on other families in the near future (he is already 
at work on the Lycaenidae). If the reader has any taxonomic interest in skippers, he/she 
must have this volume. $35.00 is cheap for the “wisdom of the ages’ in hesperiid 
taxonomy. 


LEE D. MILLER, Allyn Museum of Entomology, Florida State Museum, 3701 Bay- 
shore Road, Sarasota, Florida 33580, U.S.A. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 52 


GENERAL NOTES 


FIRST CALIFORNIA RECORD AND CONFIRMATION OF A 
ROSACEOUS HOST FOR ERIOCRANIA (ERIOCRANIIDAE) 


On 27 April 1981, numerous leafmining larvae of an eriocraniid were collected on 
Holodiscus discolor Maxim. The leafmines were abundant on Holodiscus bushes growing 
adjacent to the eucalyptus grove upslope from the entrance to San Bruno Mountain Park, 
San Mateo Co., CA. At that time, 85-90% of the mines had been abandoned by the fully 
matured larvae. 

The site was revisited on 5 February and 5 March 1982; on the second visit adults of 
Eriocrania semipurpurella pacifica Davis were found in abundance. Adults were ob- 
served from 1000 to 1200 h, during which time I observed three pairs in copulo and 
several females ovipositing on the young expanding leaves of Holodiscus. On 4 April 
1982, no adults were observed, and many of the larval mines were underway, although 
few mines had reached maturity. 

Davis (1978, Smithsonian Contrib. Zool. 251:1-131) records Holodiscus discolor as the 
probable but questionable host of E. s. pacifica based on a larval eriocraniid collection 
from Vancouver, British Columbia. All other known hosts of eriocraniids are members 
of the order Fagales. Holodiscus, a member of the Rosales, would represent a novel host 
switch for the family Eriocraniidae. The circumstantial association of larvae and adults 
and the ovipositional behavior of females confirm H. discolor as a primary host of E. s. 
pacifica. 

The geographically nearest confirmed record of this insect is Whatcom Co., WA, where 
J. F. Clarke collected adults in April 1923. However, a single male in poor condition, 
which may represent this species, was collected by Walsingham in Grant Co., OR (Davis, 
1978, loc. cit.). 

Davis (1978, loc. cit.) described E. s. pacifica as a subspecies distinct from the European 
and northeastern North American, Betula-feeding populations of E. s. semipurpurella 
(Stephens). Despite the morphological resemblance of E. s. pacifica to E. s. semipurpu- 
rella, it seems certain that the former is deserving of specific recognition as it is both 
allopatric to other known populations of E. s. semipurpurella and possesses a novel host 
association. Among leafminers it would be most unusual to find a single species feeding 
on plants belonging to different orders, i.e., the Fagales and Rosales (Needham et al., 
1928, Leaf-mining Insects. Baltimore: The Williams and Wilkens Co. 351 pp.; and Her- 
ing, 1951, Biology of Leaf Miners. s'Gravenhage: W. Junk. 420 pp.). 

Previously, three of the five described genera of Eriocraniidae were known from the 
Californian fauna: Dyseriocrania Spuler, Eriocraniella Viette, and Neocrania Davis. 
With the addition of Eriocrania semipurpurella pacifica, the Californian fauna consists 
of eight recognized species in four genera, and appears to be the richest in diversity of 
any region of comparable area. 


DAVID WAGNER, Department of Entomology, University of California, Berkeley, 
California 94720. 


VOLUME 39, NUMBER 1 53 


Journal of the Lepidopterists’ Society 
$9(1), 1985, 53 


AMPHION NESSUS (SPHINGIDAE) ATTRACTED TO PHEROMONES OF 
ANISOTA VIRGINIENSIS (SATURNIDAE) 


On the afternoon of 9 June 1983 in Groton, Middlesex County, Massachusetts, at 
approximately 1500 h, I noticed a male Amphion nessus (Cramer) hovering about an 
emergence cage containing pupae of several species of moths. It was a bright early 
summer day with temperature, humidity and wind conditions within normal ranges. 
Shortly, there were as many as four A. nessus males in the vicinity of the cage. The 
moths were searching the cage and adjacent shrubbery as though they were attempting 
to locate a “calling” female. 

The only moth in the cage was a female Anisota virginiensis virginiensis (Drury), 
whose scent organ was extended. Vegetation in the vicinity of the cage consisted of two 
evergreen shrubs and lawn. There was little possibility of a wild A. nessus female being 
in the area. Furthermore, it was obvious from the attention the cage was getting from 
the males that the A. nessus were attracted by pheromones coming from the cage itself. 

Three A. nessus males were captured easily at the cage. The anisota female was left 
in the cage for a second day and again A. nessus came to investigate. On the basis of 
visual observation, these moths were also males. 


BENJAMIN D. WILLIAMS, P.O. Box 211, Pomfret Center, Connecticut 06259. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 53-55 


A SIMPLE METHOD FOR MEASURING NECTAR EXTRACTION 
RATES IN BUTTERFLIES 


The rate at which nectarivorous animals extract nectar from flowers is one of the major 
parameters determining the instantaneous rate of energy intake, a quantity which is 
presumed to be maximized by natural selection (Pyke, Pulliam & Charnov, 1977, Quart. 
Rev. Biol. 52:187-154). The rate of energy intake equals the rate of nectar extraction 
(ul/second) multiplied by the energy content of the nectar (Joules/l). The rate of nectar 
extraction has been included in theoretical models of feeding energetics in butterflies 
(Kingsolver & Daniel, 1979, J. Theor. Biol. 76:167-179) and nectarivorous animals in 
general (Heyneman, Oecologia, 60:198-213). Although this rate has been measured in 
hummingbirds and incorporated into models of feeding energetics (Hainsworth, 1973, 
Comp. Biochem. Physiol. 46:65—78), it has apparently never been measured in butterflies 
(Kingsolver & Daniel, 1979). Here I present a simple technique for measuring extraction 
rate in butterflies which may be applicable to other nectar feeders as well. 

Nectar of a known concentration is loaded into a calibrated microcapillary tube 
(Drummond Microcaps) which is mounted on a small balsa platform with a millimeter 
scale alongside the capillary tube. The platform also includes a perch for the feeding 
butterfly to grasp (Fig. 1). The platform is displaced at a slight angle from horizontal to 
cause the nectar column to move downward as it is removed. As many butterflies main- 
tain a body temperature somewhat above ambient (Rawlins, 1980, Ecology 61:345-357), 
and since extraction rate in poikilotherms is most likely temperature dependent, I placed 
both the butterflies and the apparatus within a styrofoam chamber maintained at about 
28°C with a heat lamp. 


54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Agraulis vanillae feeding at the described apparatus. The platform encloses 
a stopwatch with which the feeding bout is timed. Alternatively, the stopwatch can be 
handheld. 


The technique for measuring extraction rate takes advantage of the apparently innate 
feeding response in butterflies which is released by the contact of the proboscis with a 
sugar solution. The butterfly is manually placed onto the perch and its proboscis is coaxed 
into contact with the leading edge of the nectar column contained in the microcapillary 
tube. As nectar extraction proceeds, the meniscus at the trailing edge of the nectar column 
can be timed with a stopwatch as it moves along the scale. I have had best success with 
100 pl microcapillary tubes (as opposed to smaller sizes), as the hardest part of the process 
is in establishing by manipulation the initial contact between the proboscis and the nectar. 
Larger microcapillary tubes have larger internal diameters, thus facilitating this part of 
the procedure. Reducing the size of the capillary tube would increase the resolution of 
the system, however. 

I have used this method with several species of papilionids, Basilarchia archippus 
(Cramer) and Agraulis vanillae (Linnaeus) (Nymphalidae) and Phoebis sennae (Lin- 
naeus) (Pieridae) with equal success. All of these species exhibit a similar response to the 
initial contact of the proboscis with the sugar solution; the proboscis begins a series of 
probing motions which sometimes pulls it out of the nectar column. If the proboscis does 
not recontact the solution within a few seconds, the butterfly coils it and ceases probing. 
If the tip is thrust back into the capillary tube, feeding begins. Once the butterfly begins 
feeding, it is no longer necessary to restrain the insect as it grasps the perch and feeds 
as it would at a flower, in some species with a characteristic folding and unfolding of 
the wings while feeding. 

Although my use of this method has been to investigate the relationship between nectar 
concentration, viscosity and extraction rate (May, in prep.), this method may also be 
useful for studies of adult diet, in which the effect of various dietary constituents on 
longevity or fecundity are measured. In studies of this type, researchers often feed the 


VOLUME 39, NUMBER 1 515) 


insects to satiation (e.g., Murphy, Launer & Ehrlich, 1983, Oecologia 56:257-263). Using 
the method described here, one can precisely control the volume of nectar imbibed by 
individual insects by regulating the volume placed within the capillary tube or by simply 
removing them from the capillary tube once a predetermined volume has been con- 
sumed. 

I would like to thank J. A. Cohen and C. S. Hieber for comments on this note. 


PETER G. May, Department of Zoology, University of Florida, Gainesville, Florida 
32611. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 55-57 


OBSERVATIONS ON THE LIFE HISTORY OF 
OCCIDRYAS ANICIA BERNADETTA (NYMPHALIDAE) 
AT THE TYPE LOCALITY 


Although Leussler’s checkerspot, Occidryas anicia bernadetta (Leussler), was de- 
scribed over 60 years ago (Leussler, 1920, Entomol. News 31:102-1083), little is known 
about its habits, and nothing has been published on the early stages or larval foodplant 
of this butterfly. Intensive collecting has been done at the type locality, Monroe Canyon 
(Sioux Co., NE); the latest report was from collections made from 1960-65 (Johnson & 
Nixon, 1967, Amer. Mid. Nat. 78(2):508-528). Even so, Leussler (1938, Entomol. News 
49:3-9, 76-80, 213-218, 275-280) sums up all that had previously been known about 
bernadetta. He states that bernadetta is “very abundant along the canyon rims in Sioux 
Co. in late May and early June.” 

In an attempt to learn more about bernadetta’s life history, two years of observations 
were made at Monroe Canyon. This report identifies a larval foodplant, describes mature 
larval and pupal stages, and identifies three parasites associated with the butterfly. 

Our experience with bernadetta began in 1982, when trips were made to Monroe 
Canyon on 22 and 29 May to search for larvae and/or adults. Several suspected foodplants 
were examined for damage, but no larvae were found. Only two adult males were seen 
and collected on 22 May. Bernadetta adults were common on 29 May, with highest 
densities observed nectaring on choke cherry, Prunus virginiana L. Adults were also 
seen resting on leaves of wolfberry, Symphoricarpos occidentalis (Hook.),. which was in 
close proximity to the P. virginiana. After watching bernadetta females alight on the S. 
occidentalis leaves, examinations of the leaves were made for ova but none were found. 
However, a pair of bernadetta were observed in copula at 1250 h, less than 0.5 m from 
the nearest S. occcidentalis plant. The pair was taken alive in an attempt to induce 
oviposition by the female, but the female died in transit. 

We returned to the type locality again on 30 May 1983, with hopes of finding im- 
mature stages of the butterfly. Chances were better for finding larvae in 1983 since the 
season was slightly retarded due to a late spring snowfall. An afternoon of collecting 
resulted in many Lepidoptera, including a few male bernadetta caught on the canyon 
slopes, but no larvae were found until the sky became overcast about 1600 h. Several 
extremely fresh male bernadetta were flushed out of the grass near a stand of S. occi- 
dentalis. A search of the S. occidentalis yielded a dozen large larvae feeding on newly 
visible leaf tips of the plants. Damage was seen only upon very close examination; it 
seemed that larval feeding was restricted to newer leaves. A thorough search of the area 
also revealed pupae and desiccated larvae. Other stands of S. occidentalis were examined 
for larvae, but only several on the higher hillsides contained larvae. Altogether, 18 larvae 
and three pupae were found on 30 May. 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


~, 
— 





Fic. 1. Larva of Occidryas anicia bernadetta on Symphoricarpos occidentalis. 


The next morning (again under an overcast sky), 27 more larvae were found, which 
led us to believe that bernadetta larvae normally fed during crepuscular to nocturnal 
hours. This might explain why larvae had been previously overlooked in the field. 

The overall color of the mature bernadetta larva is white. A thin mid-dorsal black 
stripe is interrupted by orange spots, centrally located on the dorsal area on each segment. 
A heavy, black, sub-dorsal stripe is present, and a thin, black stripe bisects the spiracular 
area. The basal color of the supraspiracular row of scoli is orange. The head and scoli 
are black, covered with black setae (Fig. 1). 

The overall color of the pupa is white. Black and orange spots and/or markings are 
present, especially on the thorax and abdomen. Wings are streaked with black. Antennal 
segments alternate black and white (Fig. 2). 

Of the 45 larvae collected, 19 (42%) were parasitized. The remaining pupated and 
eclosed as eight males and 18 females. 

The parasitized larvae were categorized by two main symptoms: (1) they would either 
shrink lengthwise, swell, and desiccate, or (2) they would remain normal size, desiccating 
only after small parasitic larvae had crawled out of the bernadetta larva and made 
cocoons nearby. The parasites that emerged from the swollen bernadetta larvae were 
identified as ichneumonids, Benjaminia sp. (probably new). Only one Benjaminia adult 
emerged per parasitized bernadetta. The parasites that emerged from the normal size 
bernadetta larvae were identified as braconids, Cotesia koebelei (Riley). Up to 30 C. 
koebelei adults emerged from a single bernadetta larva. Another ichneumonid parasite, 
Pterocormus sp., emerged from the anterior region of a single bernadetta pupa. No 
parasites were observed in the field. 

As other trips to the type locality are planned, observations on bernadetta will continue 
to be made. Other larval food plants are suspected; efforts will be made to identify them. 
The overwintering habits of bernadetta remain unknown and need to be researched. 

We wish to thank the following people for making identifications: R. C. Lommasson, 


VOLUME 39, NUMBER 1 5T 





Fic. 2. Pupae of Occidryas anicia bernadetta. 


School of Life Sciences, University of Nebraska, Lincoln, NE 68588 (S. occidentalis); V. 
K. Gupta, Center for Parasitic Hymenoptera, Gainesville, FL 32602 (Benjaminia sp., 
Pterocormus sp.); S. R. Shaw and P. M. Marsh, Systematic Entomology Laboratory, 
USDA-ABS, Insect Identification and Beneficial Insect Introduction Institute, Beltsville, 
MD 20705 (C. koebelei). 


STEPHEN M. SPOMER, Department of Entomology, University of Nebraska, Lincoln, 
Nebraska 68583 AND JAMES M. REISER, 1511 David Drive, Lincoln, Nebraska 68504. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 57-59 


NOTE ON CRUMB’ “LIBERAE ET CONFLUENTAE” 
COUPLET (NOCTUIDAE)! 


The first major systematic treatment of the larvae of North American Noctuidae was 
written by Crumb (1956, Larvae of the Phalaenidae, USDA Tech. Bull. 1185. 356 pp.). 
It is a monumental work, containing extensive diagnostic keys, larval descriptions, geo- 


1 Partially funded by the Illinois Agricultural Experiment Station Project 12-361 Biosystematics of Insects. 
2 Michigan Agricultural Experiment Station Journal Article No. 11102. 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-4. Tenth abdominal segments showing ventral and subanal regions of last 
instar noctuid larvae. 1 & 2, truncate or convex condition of posterior margin of venter 
(subanal region) (see arrows) (Alypia octomaculata); 3 & 4, medially impressed or grooved 
condition of the same region (see arrows) (Papaipema nebris). (Figs. 1 & 3 were photo- 
graphed through a Leitz Aristophot, printed sizes = 9x and 18x, respectively; 2 & 4 
were taken with the aid of a scanning electron microscope, printed sizes = 36x; all 
photographs by G.L.G.) 


graphic distributions, and a wealth of host plant information. Experienced entomologists 
as well as students taking courses on immature insects have used it with varying degrees 
of satisfaction, but many have had interpretive difficulties with the keys. 

The most obvious problems, according to a number of workers, are encountered in 
the first couplet of Crumb’s “Key to subfamilies” (p. 2) and his figures “A” and “B” in 
Plate 1. At this point users of the key encounter Crumb’s first major division of the 
noctuid larvae. He summarized the choices as “liberae” and “confluentae” in reference 
to the spatial separation between the subanal and ventral areas on the 10th abdominal 
segment. The difficulty lies, not so much in the terminology, but in the user’s trying to 
determine the perspective of the figures, which is obliquely posterior with the ventral 


VOLUME 39, NUMBER 1] 59 


side up, and in relating the lengthy couplet to the line drawings. But even when one 
knows the perspective, it is difficult to position a caterpillar in the same view under a 
dissecting microscope and still keep it submerged in alcohol. 

The purpose of our paper is to clarify this couplet by rewording it and offering light 
and SEM photographs of the appropriate structures. Hopefully, the overall utility of 
Crumb’s publication will be enhanced. In all due respect, it should be noted that Crumb’s 
publication was completed during a period of the author’s failing eyesight in his retire- 
ment years (Clarke, pers. comm.). Otherwise, we are quite certain that it would have 
been more clearly illustrated and keyed. 

Thus, our suggested alternative for the first couplet is: 


1. Venter of abdominal segment 10 not grooved posteriorly, the posterior margin 


(Sabanaleregion)-truncate or, convex (Figs. 1, 2)... 2 
Venter of abdominal segment 10 grooved posteriorly, the posterior margin (sub- 
PUKE rIOn) medially impressed (Migs. 3, 4) 6 9 


These differences show reasonably well in the accompanying illustrations of Alypia 
octomaculata Fabricius (eight-spotted forester) and Papaipema nebris (Guenée) (com- 
mon stalk borer). However, proceed with caution, because as Crumb noted, the actual 
condition is sometimes very difficult to interpret if the specimen has been inflated or has 
had its rectum everted. 


ACKNOWLEDGMENTS 


This project was aided by the advice of J. R. Byers, Entomology Research Institute, 
Agriculture Canada, Ottawa, Ontario, Canada, and H. R. Sandberg, formerly with the 
Center for Electron Microscopy, University of Illinois, Urbana-Champaign. J. F. G. Clarke, 
U.S. National Museum of Natural History, Washington, DC, is thanked for his infor- 
mation about Crumb’s career. 


G. L. GODFREY, Illinois Natural History Survey, 172 Natural Resources Building, 607 
E. Peabody Drive, Champaign, Illinois 61820, AND F. W. STEHR, Department of Ento- 
mology, Natural Science Building, Michigan State University, East Lansing, Michigan 
488 24. 


Journal of the Lepidopterists’ Society 
89(1), 1985, 59-62 


EGG PARASITISM OF APANTESIS PARTHENICE (ARCTIDAE) 
THROUGH APPARENT PHORESY BY THE WASP 
TELENOMUS SP. (SCELIONIDAE) 


On 8 Aug. 1982 while hiking along the north fork of Rock Creek near Saddlestring 
(Johnson Co.), Wyoming at the HF Bar Ranch (elev. 5400 ft) at 1400 h, I collected a 
nearly fresh female specimen of an arctiid moth, Apantesis parthenice Kirby. The insect 
was resting on sagebrush a few inches above the ground and was hand-caught. I pinched 
its thorax lightly and placed it in a folded glassine envelope (size 3.5 x 2 in) which then 
was quickly transferred into my enclosed leather collecting pouch. Upon returning to 
the ranch after several hours of hiking, all of the envelopes containing specimens taken 
that afternoon were dated and put into a plastic bag with a card containing collection 
data. This bag in turn was tightly folded, sealed with masking tape, and placed in a 
closed cigar box in a dresser drawer in my cabin at the ranch. 

Upon returning to Maryland later in the month, the cigar box was opened and un- 
packed in my laboratory at U.M.B.C. on 27 Aug. At this time I discovered 71 small dark 


60 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fics. 1-3. Arctiid moth egg shells (Apantesis parthenice and scelionid phoretic egg 
parasites, Telenomus sp.). 1, moth egg shells, showing wasp emergence holes and single 
shed puparium within each egg; 2, lateral view of wasp specimen; 3, general microscopic 
view of wasps and eggs. 


eggs which had been laid by the moth. Six first instar larvae had emerged. One of them 
was dead, but the others were crawling around inside the glassine envelope. Also found 
within the envelope were four live minute wasps (later identified as Proctotrupoidea, 
Scelionidae, Telenomus sp.). These wasps and the moth eggs are shown in Figs. 1-3. 
Further examination of all of the remaining darkened eggs using a stereo-microscope 
revealed that each contained a single fully matured wasp inside of the transparent egg 
chorion. The small wasps were living, and their wings had fully expanded within the 
eggs. The arctiid eggs all were placed into a covered plastic petri dish with a small square 
of moistened filter paper. Several wasps were watched as they emerged through the egg 
shells after chewing small holes (Fig. 1). The wasps were about 1 mm long, had yellowish 
legs and antennae, and had nearly veinless transparent round-margined wings (Fig. 2). 
By Monday 30 Aug., 52 additional wasps representing both sexes had emerged from 53 
remaining eggs (Fig. 3). Evidently, all but six of the eggs had been parasitized. 

The maternal parasitic wasp must have been introduced into the glassine envelope at 
the same time the specimen of Apantesis was placed there. There was little (if any) 
opportunity for the wasp to have found her way into the closed envelope at a later time, 
although no wasp was seen at the time of collection, and none has been found clinging 
to the pinned moth despite thorough observation through the stereo-microscope. Most 
likely the maternal wasp was clinging to the resting moth (perhaps hiding beneath her 
wings or within the thick carpet of abdominal or thoracic scales) and was inadvertently 
introduced by me into the glassine envelope in this way. 

A number of arthropods including chalcid, trichogrammatid, and scelionid wasps are 
known to exhibit phoresy, the phenomenon of attaching themselves to another insect for 
purposes of transportation (Borrer, DeLong & Triplehorn, 1964, An introduction to the 
study of insects, 4th ed., Holt, Rinehart and Winston, N.Y., 852 pp.; Comstock, 1964, An 
introduction to entomology, 9th rev. ed., Comstock Publ. Assoc., Ithaca, N.Y., 1064 pp.; 
Frost, 1959, Insect life and insect natural history, 2nd rev. ed., Dover Publ., Inc., N.Y. 
526 pp.). This behavior has been particularly well-documented for wasp species attacking 
the eggs of spiders, Hemiptera, Orthoptera, and mantids (Askew, 1971, Parasitic insects, 
Amer. Elsevier Publ. Co., Inc., N.Y. 316 pp.; Muesebeck, 1972, Nearctic species of 
Scelionidae (Hymenoptera: Proctotrupoidea) that parasitize the eggs of grasshoppers, 
Smiths. Contribs. Zool. No. 122, Smiths. Instit. Press, Wash., D.C. 33 pp.; and Rabaud, 
1922, Note sur la comportement de Rielia manticida Kieff., Proctotrupide parasite des 
ootheques de Mantes, Bull. Soc. Zool. Fr. 47:10-15). L. Van Vuuren (1935, Waarnemin- 
gen omtrent Phanurus beneficiens (Zehnt.) (Hym. Scelionidae) op Schoenobius bipunc- 
tifer Walker, Ent. Meded. Ned.-Indié 1:29-33) describes the phoretic behavior of an 
Oriental scelionid which is a common parasite of pyralid moth eggs. The female wasps 
cling to the female host (either beneath her wings or to her body) until she lays eggs, at 
which time the parasite quickly detaches herself from the host and parasitizes the freshly 
laid eggs. This relationship is not an obligatory one, however. 


VOLUME 39, NUMBER 1 61 


TABLE 1. Records of (A) Telenomus wasps (Scelionidae) known to attack arctiids 
(with known localities of occurrence) and (B) parasitoids having Apantesis moth hosts, 
based on Muesebeck et al., 1951 and Krombein et al., 1979 (information courtesy of R. 
T. Mitchell). 


Moth hosts Wasp parasites 


A) Telenomus species known to attack arctiids: 


Hyphantria cunea (Drury) Telenomus bifidis Riley, D.C., Mo. 

Diacrisia virginica (Fabr.) T. nigriscapus Ashm., Mich., IIl. 

Diacrisia virginica (Fabr.) T. spilosomatus Ashm., D.C., Va., Kans. 
B) Parasitoids having Apantesis moth hosts: 

Apantesis mais (Drury) Coelopisthia forbesii (D.T.) Pteromalidae 

A. virgo (L.) Casinaria genuina (Nort.) Ichneumonidae 

Apantesis sp. Hyposoter rivalis (Cress.) Ichneumonidae 

Apantesis sp. Apanteles phobetri Rohwer, Brachonidae 


Since the A. parthenice moth’s thorax had been pinched, she probably oviposited over 
the next day or two only following collection (9-10 Aug. 1982). The wasp parasite in 
turn must have laid her own eggs at that time (or shortly thereafter) as well. Both moth 
egg-laying and wasp egg location and parasitization within the confines of the glassine 
envelope must have occurred in totally dark conditions inside the closed cigar box and 
closed dresser drawer. Development time of the Apantesis larvae and the full maturation 
of the proctotrupoid wasps both required between 17-19 days, basically at room tem- 
perature (i.e., indoors). 

Whether or not these tiny egg parasites of Lepidoptera often search for, locate, and 
remain with females of their hosts, as these observations suggest, needs to be verified in 
the field. Such an egg-locating strategy would seem to be a very efficient one, especially, 
if fresh soft eggs are required for successful parasite oviposition. The overall frequency 
of moth egg parasitism was 65/71 eggs (or 91.5%) within the cramped confines of the 
glassine envelope. Such phoretic behavior would seem to convey a tremendous selective 
advantage to those individuals which practice it, as compared to the alternative strategy 
of directly seeking out individual eggs (or egg masses in the case of A. parthenice). Since 
only a single wasp parasite emerged from each moth egg, it is likely the maternal wasp 
was capable of distinguishing unparasitized (e.g., newly laid eggs) from those she had 
previously parasitized. Such chemosensory capabilities are widely known among parasitic 
Hymenoptera. 

Although numerous parasitoids of the arctiidae are known, neither Muesebeck et al. 
(1951, Hymenoptera of America north of Mexico. Synoptic Catalogue, U.S. Govt. Print. 
Off., Wash., D.C. 1420 pp.) nor Krombein et al. (1979, Catalogue of Hymenoptera in 
America north of Mexico, Smiths. Inst. Press, Wash., D.C. Vols. 1 and 2, 275 pp.) list 
any parasitoids for A. parthenice. Only six wasp genera are included for the entire moth 
genus, as shown in Table 1. Fifty-three species of Telenomus wasps have been described 
from North America, but only three of them (as listed in the Table) are known to attack 
arctiid moths. The species of Telenomus here described may possibly be a new one, 
since this host record is new, and those previously reported are either eastern or mid- 
western. Also, only about one quarter of the total telenomid species in North America 
have so far been described (P. M. March, pers. comm.). This possibility presently is being 
investigated further. Scelionid wasps in some cases have been successfully used as bio- 
logical control agents for insect pests. 

I am grateful to Dr. P. M. Marsh for identifying the wasps, and to Dr. D. C. Ferguson 
for confirming the moth species. Both persons are from the Systematic Entomology 
Laboratory, U.S.D.A., U.S. Natural History Museum, Wash., D.C. 20560. I thank R. T. 
Mitchell of Silver Spring, Maryland for providing the information contained in Table 1, 
and for comments on the manuscript. 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Covell, Jr. (1984. A field guide to the moths of eastern North America. Houghton 
Mifflin Co., Boston. 496 pp.) places this tiger moth in genus Grammia. Dr. Norman 
Johnson, Department of Entomology, Ohio State University, presently is revising the 
taxonomy of Scelionid wasps. He recently informed me (pers. comm.) that most of the 
early type specimens of Telenomus are females but that the male genitalia possess 
important diagnostic features for determining species status. This wasp species in his 
opinion may be undescribed and no specific designation can be given at this time. 


AUSTIN P. PLATT, Department of Biological Sciences, University of Maryland Bal- 
timore County, 5401 Wilkens Avenue, Catonsville, Maryland 21228. 


Journal of the Lepidopterists’ Society 
39(1), 1985, 62-63 


NOTES ON THE HABITAT AND FOODPLANT OF INCISALIA HENRICI 
(LYCAENIDAE) AND PYGRUS CENTAUREAE (HESPERIIDAE) 
IN MICHIGAN 


The foodplant of Incisalia henrici (Grote and Robinson) in Michigan was unknown 
until 1981, when it was confirmed that maple-leaf viburnum, Viburnum acerifolium L. 
(Caprifoliaceae) is an acceptable foodplant. According to Tietz (1972, An index to the 
described life histories, early stages, and hosts of the Macrolepidoptera of the Continental 
United States and Canada, Allyn Mus. Entomol., Sarasota, FL) and Pyle (1981, The 
Audubon Society field guide to North American butterflies, A. Knopf, Inc., NY), vibur- 
num is not listed as a known foodplant for I. henrici. 

I first became acquainted with Henry’s Elfin in 1953, when a series was collected in 
the Langston State Game Area, Montcalm County, on 15 and 23 May. Since that time, 
I. henrici has been collected and observed in the same area in close proximity to second 
growth aspen (Populus grandidentata Michx. and tremuloides Michx.), white oak (Quer- 
cus alba L.) and red maple (Acer rubrum L.), with scattered white pine (Pinus strobus 
L.) on sandy soil. Most of the adults have been taken (before full leaf development along 
sandy trails and narrow wooded sunny openings) while perched on small shrubs, on dried 
leaves and twigs or on bare sand. At this site, adults could easily be overlooked because 
of their small size and dark color. Only once was an adult observed nectaring on choke 
cherry, Prunus virginiana L., along the trail. During this period, the elfin gave no clues 
to the preferred larval foodplant despite the presence in the Game Area of Prunus sp. 
and Vaccinium sp., two previously recorded foodplants for I. henrici. 

It wasn’t until 3 June 1979, that Harry King and I discovered several Lycaenidae 
larvae feeding on the flower cymes of V. acerifolium in a similar aspen-oak woods, 
located one and one-half miles north of the original site. The greenish slug-shaped larvae, 
with pale lateral stripes, appeared to resemble I. henrici, based on the brief description 
in Klots (1951, Field guide to the butterflies, Houghton Mifflin Co., MA). The larvae 
were removed and kept in captivity until the following spring when (to my disappoint- 
ment) Celastrina ladon (Cramer) emerged. Then during 1980-1982, I examined flower 
cymes of V. acerifolium at both Game Area locations and found numerous larvae of 
various instars representing C. ladon and what was believed to be I. henrici. Subsequent 
emergence of I. henrici in 1981 and 1983 from over-wintering pupae finally confirmed 
the use of Viburnum acerifolium as the preferred foodplant in this location. 

In 1974, Larry West, noted nature photographer, observed a female Pygrus centaureae 
wyandot (Edwards) oviposit an egg on the underside of a wild strawberry leaf, Fragaria 
virginiana Duchesne, on 22 May in Otsego County, Michigan. Since 1958, the grizzled 
skipper has been collected from 15 May to 3 June on a pine barren in an area of short 


VOLUME 39, NUMBER 1 63 


grasses and sedges (including Danthonia spicata (L.) Beauv. and Carex pennsylvanica 
Lam.) on sandy soil. This skipper is not easily seen on the wing but can be collected with 
some frequency while nectaring on wild strawberry scattered in large patches throughout 
the open areas. Butterfly species that occur in the same area during the approximate 
flight period of P. centaureae include Euchloe olympia (Edwards), Oeneis chryxus stri- 
gulosa McDunnough and Hesperia metea Scudder. 

With wild strawberry as the possible foodplant for P. centaureae, I searched strawberry 
patches during the summer from 1975 to 1979 for signs of larvae. Several mid-instar 
larvae were finally found in leaf nests on wild strawberry; the nests varied from a single 
folded leaf to three leaves held together with silk. The larval nests were constructed so 
the larvae rested on the upper leaf surface. Frequently, the heat of the day would curl 
many leaves, or a spider would curl a leaf for its egg mass, making it frustrating and 
difficult to find P. centaureae larval nests. The larvae were removed to captivity and 
finished feeding by late summer and over-wintered in the pupa stage. In reviewing the 
literature, this is the first record of wild strawberry as the foodplant for P. centaureae 
wyandot; other authors (Pyle, ibid.; and Ferris & Brown, 1981, Butterflies of the Rocky 
Mountain states, Univ. Oklahoma Press, OK) have cited Rubus and Potentilla (Rosaceae) 
as foodplants for P. centaureae in other parts of its range. 

Perhaps both species will prove to be more widespread in the Great Lakes region when 
collectors are aware of their habitat and foodplant requirements. I wish to express my 
deep appreciation to Harry King and Larry West for sharing their field observations 
with me. 


MOGENS C. NIELSEN, Adjunct Curator, Department of Entomology, Michigan State 
University, East Lansing, Michigan 48824. 


Date of Issue (Vol. 39, No. 1): 16 October 1985 


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EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


_IsA MONTENEGRO, Editorial Assistant 
DouGLas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 
NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
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heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Ady. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

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CONTENTS 


New U.S. RECORDS AND OTHER INTERESTING MOTHS FROM 
TExAs. André Blanchard ¢y Edward C. Knudson ........... 
NOTES ON THE LARVA AND BIOLOGY OF MOODNA BISINUELLA 
HAMPSON (PYRALIDAE: PHYCITINAE). H. H. Neunzig y 
BIOLOGY AND DESCRIPTION OF THE LARVA OF DICYMOLOMIA 
METALLIFERALIS: A CASE-BEARING GLAPHYRIINE (PYRALIDAE). 


David Wagner oe 
FIELD SURVEY OF THE TRUE BUTTERFLIES (PAPILIONOIDEA) OF 
RHODE ISLAND: Harry Pavulaan 0. 


FOREST TORTRICIDS TRAPPED USING EUCOSMA AND RHYACIONIA 
SYNTHETIC SEX ATTRACTANTS. Ri. E. Stevens, C. Sartwell, 
T. W. Koerber, J. A. Powell, G. E. Daterman & L. L. Sower 
NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF OPSI- 
PHANES QUITERIA QUIRINUS GODMAN AND ERYPHANIS AE- 
SACUS BUBOCULUS BUTLER (BRASSOLIDAE). Rolando Cub- 


AKO ALL aA NA NA Ae MEL MMO SEMEN ONGIALAITRRMO ED OAL TN ee ee 
THE DEFENSIVE ENSEMBLES OF TWO PALATABLE Motus. David 
Di BO0ans) es ee 
CYMAENES FINCA, SP. N. (HESPERIIDAE) FROM TRINIDAD, W.I. 
M. J. Wi: Cocke 


GENERAL NOTES 


First California record and confirmation of a rosaceous host for Eriocrania 
(Eriocrantidae). David Wagner, 2 


Amphion nessus (Sphingidae) attracted to pheromones of Anisota virginiensis 
(Saturniidae). Benjamin D. Williams 00 ea 


A simple method for measuring nectar extraction rates in butterflies. Peter 
Ge. Mag oi ee a 


Observations on the life history of Occidryas anicia bernadetta (Nymphalidae) 
at the type locality. Stephen M. Spomer & James M. Reiser 0. 


Note on Crumb’s “liberae et confluentae” couplet (Noctuidae). G. L. God- 
predids BOW Stehr vii ik ol I 


Egg parasitism of Apantesis parthenice (Arctiidae) through apparent phoresy 
by the wasp Telenomus sp. (Scelionidae). Austin P. Platt cco 
Notes on the habitat and foodplant of Incisalia henrici (Lycaenidae) and 
Pygrus centaureae (Hesperiidae) in Michigan. Mogens C. Nielsen. ...... 


BOOK’ REVIEW i an 


9 


62 


eo 


——— 


Volume 39 1985 Number 2 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





7 January 1986 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Don R. Davis, President LEE D. MILLER, 

ViTorR O. BECKER, Vice President Immediate Past President 
JAVIER DE LA Maza E., Vice President JULIAN P. DONAHUE, Secretary 
JOHN C. DOWNEY, Vice President Eric H. METZLER, Treasurer 


Members at large: 


F. S. CHEW J. M. BuRNS B. A. DRUMMOND 
G. J. HARJES F. W. PRESTON J. LANE 
E. H. METZLER N. E. STAMP R. K. ROBBINS 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
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itate the exchange of specimens and ideas by both the professional worker and the 
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Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
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Cover illustration: Micropylar end view (130) of the egg of Sericosema sp. (probably 
juturnaria) (Geometridae). The scanning electronmicrograph was taken by Thomas D. 
Eichlin, Sacramento, of eggs furnished by Ron Robertson, Santa Rosa, California. 


JoURNAL OF 
Tue LeEerPipopreRIstTs’ SOCIETY 


Volume 39 1985 Number 2 


Journal of the Lepidopterists’ Society 
89(2), 1985, 65-84 


MAINTAINING SPECIES INTEGRITY BETWEEN 
SYMPATRIC POPULATIONS OF HYALOPHORA CECROPIA 
AND HYALOPHORA COLUMBIA (SATURNIIDAE) 

IN CENTRAL MICHIGAN 


JAMES P. TUTTLE 
728 Coachman #4, Troy, Michigan 48083 


ABSTRACT. The available literature suggests that Hyalophora cecropia and Hya- 
lophora columbia have identical ethological requirements with respect to reproductive 
behavior and F, hybrids have often been produced in the laboratory. As a result, investi- 
gators have long been puzzled by the rarity of natural hybrids since the two species 
occur sympatrically over much of the northern Great Lakes region. 

Field observations in Montcalm County, Michigan, identified a series of progressive 
barriers which operate to prevent hybridization. The prezygotic isolating mechanisms, 
seasonal isolation, daily allochronic flight behavior, and mechanical incompatibility re- 
strict most of the interaction between the two species. However, when individuals oc- 
casionally overcome those barriers and a successful hybrid mating occurs, several post- 
zygotic isolating mechanisms are then tested. As a result of the overall effectiveness of 
these systems, the gene pool of each species is protected, thereby allowing the two species 
to co-exist. 


The large moths of the genus Hyalophora (Saturniidae) are repre- 
sented by two species within the state of Michigan. The cecropia moth, 
Hyalophora cecropia (L.), is widely distributed, while Hyalophora co- 
lumbia (S. I. Smith) is a much more localized species. 

When Moore (1955) published his checklist of the Michigan moths, 
he listed 18 county records for H. cecropia from such varied locales as 
Dickinson County in the western portion of the Upper Peninsula to 
metropolitan Wayne County in the southeastern portion of the Lower 
Peninsula. He also cited statewide adult capture dates ranging from 
27 May to 5 August.! The author’s recent review of additional material 
in public and private collections has increased to 48 the number of 
counties from which H. cecropia has been reported (Fig. 1). The pa- 


The 5 August date is based upon a reared specimen probably maintained under artificial conditions. Such late dates 
are highly unlikely in nature. 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





COLUMBIA | ¢ | 


CECROPIA 





Fic. 1. Distributional map of H. cecropia and H. columbia in Michigan. Montcalm 
County (darkened) is the site of the present study. 


rameters of the adult flight period were also expanded when wild 
capture dates as early as 22 May were discovered. 

The distribution of H. cecropia within the state is greatly enhanced 
by the acceptance of a wide array of larval foodplants. Tietz (1972) 
lists 75 specific larval foodplant records, which include many of the 
native Michigan broadleaved trees and shrubs. This adaptability to 
varied plant communities is extremely important to a cosmopolitan 
species such as H. cecropia. 

Although there are still a number of counties from which H. cecropia 
has not been reported, this would appear to be due to a lack of col- 
lecting. Based upon the adaptability of the species, the lack of restric- 


VOLUME 39, NUMBER 2 67 


tive topographical barriers within the state and the distribution of ex- 
isting county records, it is reasonable to assume that H. cecropia occurs 
in all of Michigan’s 83 counties. 

In contrast to the large amount of data available on H. cecropia, the 
existing Michigan records of the distribution and flight period of H. 
columbia are extremely limited. Based exclusively on wild collected 
cocoons, Moore (1955) cited records for H. columbia from Lapeer, 
Montcalm, Oakland, Washtenaw, and Wayne counties. Since the pub- 
lication of Moore’s checklist there have been records from Ingham 
(Collins & Weast, 1961), Dickinson, Jackson, Livingston, Mecosta, Ne- 
waygo (Ferguson, 1972), Clare, Menominee, Schoolcraft (M. C. Niel- 
sen, in litt.), and Isabella counties (Ted Herig, pers. comm.) (Fig. 1). 
Most of these additional records are also based upon wild collected 
cocoons. 

In fact, prior to the present study, there were only six reports of H. 
columbia adults having been collected in Michigan. Three adults have 
been collected at light: a male collected in Menominee County on 10 
June 1971; a female collected in Schoolcraft County on 21 June 1971; 
and a male (specimen lost) collected in Dickinson County (M. C. Niel- 
sen, in litt.). In addition, in June 1978, single H. columbia males were 
attracted on each of three successive nights to a H. cecropia female in 
Montcalm County (Frank Hedges, pers. comm.). 

Unlike the widespread distribution of H. cecropia, the range of H. 
columbia is limited by its dependency on larch (Larix laricina) as the 
larval foodplant. Although there are records of H. columbia being 
reared in captivity on choke cherry (Prunus virginiana) (Collins & 
Weast, 1961) and weeping willow (Salix babylonica) (Norman Trem- 
blay, pers. comm.), it is now generally accepted that the larvae feed 
exclusively on larch in nature. 

Many old foodplant records were erroneously based on cocoons found 
attached to plants growing in association with larch. This is clearly 
reflected in the type description of H. columbia, when Smith (1863) 
cites ““Nemopanthes canadensis and Rhodora canadensis” as larval 
foodplants and only incidentally mentions larch as a possible alternate 
host. This can now be explained by the tendency of full-fed larvae to 
wander in search of an appropriate cocoon spinning site. 

In Michigan’s Upper Peninsula larch abounds as a dominant tree 
species in conifer bog and swamp community associations (Barnes & 
Wagner, 1981). As a result of the widespread distribution of larch in 
this area, H. columbia should be expected to occur regularly. The 
relatively few records from this region are almost certainly due to 
limited collecting. 

The existing county records for H. columbia in Michigan’s Lower 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Peninsula occur as two separate pockets which include seven counties 
in the southeast and five counties in the central portion of the penin- 
sula. In these areas, larch occurs only in poorly drained “kettle hole” 
type situations. Such bogs may be isolated by several miles of unsuitable 
habitat and are historically very susceptible to human influence. 

An examination of specimens at the University of Michigan reveals 
several records for H. columbia from Wayne County prior to 1900. 
However, drainage and the resulting development have eliminated all 
of the suitable habitat. As a result, there are no records of H. columbia 
from Wayne County in this century. 

H. columbia populations in Michigan’s Lower Peninsula are, there- 
fore, restricted to those undisturbed areas where larch occurs in suffi- 
cient numbers to effectively support the insect. One such locale is the 
Stanton State Game Area in Montcalm County (Fig. 1). In this area, 
which serves as the site of the present study, H. columbia occurs sym- 
patrically with H. cecropia. 

Within zones of contact between closely related species such as H. 
cecropia and H. columbia, effective mate recognition behavior must 
be established if both species are to co-exist over an extended period 
of time (Mayr, 1970). Yet, the available literature suggests that H. 
cecropia and H. columbia lack such discriminatory behavior. 

The existing data indicate that H. cecropia and H. columbia adults 
emerge at the same time of the year (Moore, 1955; Collins & Weast, 
1961; Ferguson, 1972). Both species also mate during the hours im- 
mediately preceding dawn (Collins & Weast, 1961; Ferguson, 1972). 

In addition, even the earliest breeders were able to obtain interspe- 
cific Hyalophora hybrids in the laboratory (Morton, 1895; Soule, 1907). 
Since that time, almost all possible crosses, backcrosses, and reciprocal 
crosses have been attempted (Sweadner, 1937; Weast, 1959; Collins & 
Weast, 1961; Wright, 1971; Kohalmi & Moens, 1975). 

Investigators thus learned that congeneric males would readily re- 
spond to calling” non-specific females. Sweadner (1937) effectively ex- 
ploited this ability by using H. cecropia females to attract wild H. 
columbia and H. gloveri nokomis (Brodie) males in western Ontario 
and Manitoba. More recently, H. cecropia females have been success- 
fully used to attract wild H. columbia males in Manitoba (Collins, 
1973), Ontario (Kohalmi & Moens, 1975), Quebec (Gilles Deslisle, pers. 
comm.), Wisconsin (Ferge, 1983), and Michigan (Frank Hedges, pers. 
comm.). 

Yet, in spite of these apparent behavioral similarities and the relative 
ease with which hybridization occurs in captivity, very few valid nat- 


* Extension of the ovipositor resulting in the release of a sexual pheromone. 


VOLUME 39, NUMBER 2 69 


ural hybrids between H. cecropia and H. columbia have been reported. 
The author agrees with Ferguson (1972) who states, “Careful scrutiny 
of the literature reveals that at least some of the records of natural 
hybrids may be false, especially those based on cocoons only. Free- 
man’s experience of having a normal cecropia emerge from what looked 
like a hybrid cocoon on larch, already noted, casts suspicion on all 
reports based on cocoons of intermediate appearance.” 

Ferguson (1972) does cite two records of hybrids emerging from 
cocoons collected in Mecosta County, Michigan, and Fraserburg, On- 
tario. He also mentions a wild collected hybrid male collected on 18 
June in Oakland County, Michigan. The author has examined a wild 
male collected on 17 June 1973, in Oakland County, Michigan, which 
appears to reflect a hybrid background. During the present study in 
Montcalm County, only one of 259 Hyalophora adults examined showed 
any hybrid influence. 

In the spring of 1981, the author observed the emergence of Hya- 
lophora adults in outdoor cages in Ann Arbor, Michigan. An extended 
emergence pattern was observed in the caged H. cecropia adults. A 
comparison with the concentrated emergence of the H. columbia adults 
in the adjacent cage suggested partial temporal isolation. These obser- 
vations supplied the impetus for the present study which offers data 
and discusses the reproductive isolating mechanisms which allow such 
closely related species to co-exist. 


MATERIALS AND METHODS 


Field studies were conducted at two sites in the immediate vicinity 
of the village of Stanton in Montcalm County. The two bogs are ap- 
proximately 3 km apart and separated by farmland and the small 
Stanton residential community. 

Throughout this area, naturally occurring and ornamentally planted 
maple (Acer), oak (Quercus), elm (Ulmus), and aspen (Populus) pre- 
dominate. Within the bogs, which are dominated by larch, there also 
occur willow (Salix), cherry (Prunus), and the lower growing dog- 
woods (Cornus). 

At the first Stanton site a five foot tall trap, modeled after the type 
proven effective by Sternburg and Waldbauer (1969), was erected near 
the edge of the bog. The trap was situated so that pheromone released 
by bait H. cecropia females was carried throughout the bog by pre- 
vailing westerly winds. Trap construction did not allow responding 
wild males access to the confined females. As a result, the males are 
drawn into the trap and eventually settle on the sides. The males were 
collected daily, killed and then stored for later examination. 

In order to maintain an adequate supply of bait females for the trap, 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


approximately 300 H. cecropia cocoons’ were taken from the outdoors 
on 1 January of 1982 and 1983 and placed in a temperature controlled 
environment of 6°C. Starting in the middle of March of each year, 
cocoons were removed from the controlled environment, slit open, and 
sexed by examining the pupal antennae (Miller, Highfill & Cooper, 
1982) until a female was identified. Any male pupae were discarded 
and the female pupa was placed in an emergence chamber at room 
temperature (20°C). From that date forward and until the first week 
of June, at least one additional female was placed in the emergence 
chamber each day. 

Within approximately six weeks the bait females began emerging. 
They were placed in the trap on the day of their emergence and were 
not removed until after they had died. These unmated females lived 
5-10 days and continued to call each morning from approximately 
0330 h (EDST) until well after sunrise. 

By maintaining the above described schedule, the trap contained at 
least one, and usually several, calling H. cecropia females every day 
from 3 May-15 July 1982. The following year, the trap contained at 
least one calling H. cecropia female every day from 1 May-20 July 
1988. The duration of availability of calling females in the trap ex- 
tended well beyond the anticipated emergence season of the local H. 
cecropia and H. columbia populations. Therefore, the wild Hya- 
lophora males responding to the trap reflect the natural seasonal flight 
periods of both species. 

Since the bait-trap samples only males, a second method was also 
used to gather data on the seasonal flight periods of both species. Dur- 
ing the fall of 1981 and 1982, H. cecropia and H. columbia cocoons 
reared from Stanton area stock were stored in wire cages near the 
village of Sheridan (approximately 9 km from the Stanton sites) and 
exposed to natural weather conditions. During the spring of 1982 and 
1983, the emergence date and sex of each eclosing adult in this caged 
group were recorded, and the resulting data were compared with the 
trap data. 

At the second Stanton site, the H. cecropia and H. columbia females 
emerging from the caged group kept at the Sheridan site served an 
additional purpose. Females of both species were alternately tied-out* 
at 50 m intervals along the western edge of the bog. Unlike the bait 
females in the trap at the first site, the females at this site were readily 
accessible to responding males. These females were checked at 15 mi- 


5 Livestock was reared by the author and also obtained from other breeders. 
* Tethering of an unmated female to a tree, shrub, or other fixed object. 


VOLUME 39, NUMBER 2 71 


TABLE 1. Isolating mechanisms between H. cecropia and H. columbia in Montcalm 
County, Michigan. 








I. Precontact Mechanisms 
A. Seasonal isolation 
1. species variation in annual availability of adults 
2. variation in male-female emergence 
B. Daily allochronic flight behavior 


II. Contact Mechanisms 
A. Mechanical isolation 
B. Intraspecific remating 


III. Postmating Mechanisms 
A. Embryonic mortality 
B. Larval acceptance of oviposition substrate 
C. Inviability of F, progeny 
1. larval 
2. pupal 
3. adult 
D. Hybrid sterility* 
E. Activity of hybrid adults* 
1. seasonal emergence 
2. daily flight behavior 
Note: Precontact Mechanisms (I) and Contact Mechanisms (II) are prezygotic, the transfer of gametes not having 
taken place. Bosman ng Mechanisms (III) are postzygotic, hybrid zygotes ae been formed. 
Due to the limited number of hybrid adults obtained, these factors could not be addressed but are offered to 
complete the table. 


nute intervals from sunset until 0300 h and were then continuously 
monitored until well after sunrise to determine when each female 
began calling. Males visually identified as conspecific with the female 
to which it was responding were netted and held for later examination; 
the female being allowed to continue calling. Males involved in inter- 
specific contacts were allowed to attempt copulation. Only those males 
which began to “hover” within 1-3 m of a calling female or actually 
attempted copulation were scored in the response count. 

In addition, during the course of the study, a limited number of 
pairings between H. cecropia and H. columbia were obtained. In each 
case, the activities of the mated females were closely monitored, mor- 
tality during all developmental stages was recorded, and the behavior 
of the hybrids in all stages was noted. 


RESULTS 


Table 1 lists the isolating mechanisms between H. cecropia and H. 
columbia in Montcalm County, Michigan. The table is broken down 
into three sequentially applied categories: Precontact Mechanisms, 
Contact Mechanisms, and Postmating Mechanisms. 

The first Precontact Mechanism is seasonal isolation which occurs in 
two facets. The primary facet of seasonal isolation involves variation 
in the annual emergence pattern of each species. 


2; JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





15 — 

14 — 

13 — g 

12 — E (2A) 

ie 

10 — Columbia (41) 
gee Cecropia (65) Hi 


NUMBER OF 
MALES CAPTURED 








10 — 
9 — 
8 — (28) 
7- Columbia (15) 
6 Cecropia (35) Hi 


NUMBER OF 
ADULT EMERGENCES 
oO 
| 





MAY JUNE 


(3A) 


Columbia (33) 
Cecropia (62) Hi 


NUMBER OF 
MALES CAPTURED 


— 
= wo & Om 4 wo wo O 





10 — 
ae (3B) 
3 E = Columbia (15) 7 
33 A ne Cecropia (38) Hi 
P65 
3 4 
3 
2 
1 





JUNE JULY 


Fics. 2-3. Comparative data for H. cecropia and H. columbia. 2A, trap capture of 
wild males at the first Stanton, Michigan site in 1982. 2B, total cage emergence of adults 
at Sheridan, Michigan in 1982. 3A, trap capture of wild males at the first Stanton, 
Michigan site in 1983. 3B, total cage emergence of adults at Sheridan, Michigan in 1983. 


VOLUME 39, NUMBER 2 73 


A total of 65 wild H. cecropia males was attracted to the trap from 
27 May-7 July 1982 (Fig. 2A). The trap data indicate an early phase 
of the H. cecropia flight period occurred from 27 May-5 June 1982. 
This phase involved 9 males or 13.8% of the total H. cecropia trap 
sample. The main phase of the H. cecropia flight period occurred from 
17 June-7 July 1982. 

A total of 41 wild H. columbia males was attracted to the trap from 
28 May-9 June 1982 (Fig. 2A). In contrast to the extended flight period 
of H. cecropia, the range of dates for H. columbia is very concentrated. 

The emergence pattern of the 1982 caged group at Sheridan, Mich- 
igan (Fig. 2B) compares quite favorably with the trap results. There is 
a shift to the right in the trap capture dates (Fig. 2A) when compared 
to the emergence dates of the caged group (Fig. 2B). However, as 
pointed out by Sternburg and Waldbauer (1969), such a shift is to be 
expected. 

During the replicate field studies of 1988, a total of 62 H. cecropia 
males was collected in the trap between 10 June-14 July 19838 (Fig. 
3A). The early phase of the H. cecropia flight period extended from 
10-17 June 1983 and involved only four males or 6.4% of the total H. 
cecropia trap sample. The main phase of the H. cecropia flight period 
occurred from 26 June-14 July 1983. 

A total of 83 wild H. columbia males was captured in the trap 
between 14-22 June 1983 (Fig. 3A). As in the previous year, the H. 
columbia flight period is very concentrated. 

Figure 3B depicts the emergence of the 1983 caged group at Sher- 
idan, Michigan. As in the preceding year, the emergence pattern of 
the caged group (Fig. 3B) parallels the trap captures (Fig. 3A). 

The second facet of seasonal isolation involves variation in male- 
female emergence. Such an analysis was based on the emergence pat- 
tern of the H. columbia and those early phase H. cecropia in the caged 
groups at Sheridan, Michigan (Figs. 2B & 3B). 

In the 1982 caged group (Fig. 2C), the potential for contact between 
H. columbia males and H. cecropia females clearly existed, since all 
emergences occurred between 24-29 May 1982. However, the poten- 
tial for reciprocal contact was greatly reduced, since the H. cecropia 
males emerged from 20-24 May 1982, and the H. columbia females 
emerged from 28 May-2 June 1982. 

In the 1983 caged group (Fig. 3C), a similar pattern was observed 
when H. columbia males and H. cecropia females emerged between 
13-18 June 1983. The H. cecropia males emerged on 9-10 June 1983, 
while the H. columbia females emerged between 17-22 June 1983. 

The second type of Precontact Mechanism is daily allochronic flight 
behavior. Hyalophora females tied-out at the second Stanton site were 


+] 
aN 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


—_ 
© 
| 


(2C) 


Columbia & WN 
BS 


a ~~ co o 
a aie 





NUMBER OF ADULT EMERGENCES 


(3C) 
10 — Columbia oN 
g — Q Be 
8 — Cecropia Oe 


i or 


NUMBER OF ADULT EMERGENCES 





JUNE 


Fics. 2C & 3C. The cage emergence of early phase H. cecropia and H. columbia 
adults at Sheridan, Michigan. 2C, in 1982. 3C, in 1983. 


VOLUME 39, NUMBER 2 To 


closely monitored during the two year study. The earliest time a H. 
cecropia female was observed calling was 0325 h and unmated females 
continued to call for almost two hours after sunrise. During the same 
periods, the earliest time a H. columbia female was observed calling 
was 0445 h. As with H. cecropia, unmated H. columbia females con- 
tinued to call well after sunrise. 

Figure 4 depicts the arrival time of wild Hyalophora males to calling 
Hyalophora females at the second Stanton site in 1982. The data pre- 
sented are the results of pooled observations of male arrivals from 29 
May-1 June 1982. The first H. cecropia male arrived at 0335 h and 
the uninterrupted response continued until 0450 h. Only two H. ce- 
cropia males were attracted after sunrise, which occurred at 0500 h. 
In contrast, the wild H. columbia males did not begin responding until 
after daybreak. The first H. columbia male was attracted at 0520 h, 
and males continued to respond until 0625 h. 

Figure 5 illustrates the arrival time of wild Hyalophora males to 
calling Hyalophora females at the second Stanton site in 1983. Pooled 
observations of male arrivals were made from 17-19 June 1983. The 
lone H. cecropia male taken during the year was attracted at 0445 h. 
One H. columbia male was attracted at 0435 h, which was the only 
record of a H. columbia male responding prior to sunrise (0450 h). 
The remainder of the H. columbia response began at 0508 h and 
continued until 0614 h. 

As a result of maintaining a continuous release of pheromone and 
the apparent lack of pheromone differentiation, a number of interspe- 
cific contacts were noted at the second Stanton site. During 1982 (Fig. 
4), 14 conspecific contacts were scored: 8 H. cecropia contacts and 6 
H. columbia contacts. During that same time period, 16 interspecific 
contacts were scored, all of those contacts being H. columbia 6 x H. 
cecropia 2. During 1988 at the second Stanton site (Fig. 5), 23 conspe- 
cific contacts were scored, all of those being H. columbia contacts. 
During that same time period five interspecific contacts were scored: 
4 H. columbia é x H. cecropia 2 and 1 H. cecropia 8 x H. colum- 
bia 2.5 

During the two year study only one contact between a H. cecropia 
male and a H. columbia female was observed. On 17 June 1988, at 
0445 h, a male H. cecropia coupled with a calling H. columbia female 
at the second Stanton site. As a result of the pairing, the genitalia of 
the female was ruptured, and she soon died. Previous investigators had 


5On 19 June 1983, only calling H. columbia females were available and 11 conspecific H. columbia contacts were 
scored. As a result, the total ratio of conspecific to interspecific contacts was significantly higher in 1983. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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TIME OF MALE ARRIVALS (E.D.S.T.) 
Fics. 4 & 5. The arrival times of wild H. cecropia and H. columbia males to calling 


Hyalophora females at the second Stanton, Michigan, site. 4, data based on pooled 
observations from 29 May-1 June 1982. 5, data based on pooled observations from 17-— 


19 June 1983. 


VOLUME 39, NUMBER 2 rig 


similar experiences while attempting crosses between the Callosamia 
(Peigler, 1977) and other Hyalophora (Collins & Weast, 1961). 

In contrast, 20 contacts between H. columbia males and H. cecropia 
females at the second Stanton site were scored during the study. On 
most every occasion the author observed the H. columbia male make 
numerous unsuccessful attempts to clasp onto the larger calling H. 
cecropia female and then fly away. 

On only two occasions did the author see a H. columbia male “‘suc- 
cessfully” clasp onto a calling H. cecropia female. On 1 June 1982, at 
0608 h, a wild H. columbia male coupled with a H. cecropia female. 
The female appeared uneasy during the entire contact and after 15- 
20 minutes managed to break away from the male. On 2 June 1982, 
the same female was mated to a wild H. cecropia male in Troy, Mich- 
igan. The resulting offspring were reared through as pure H. cecropia. 

Even more interesting were observations of a wild H. columbia male 
coupled with a H. cecropia female at 0485 h on 17 June 1983. The 
pair remained in copula until 0910 h. At dusk on that same day the 
H. cecropia female immediately laid 44 ova and then settled on the 
side of the cage. 

On 18 June 1983, at 0345 h, the same H. cecropia female once again 
began calling. At 0522 h the female attracted another H. columbia 
male but was not allowed to mate. During that evening the female 
made no attempt to lay any additional ova. 

On 19 June 1983, the same H. cecropia female was cage-mated to 
a H. cecropia male from Troy, Michigan. During that evening 161 
additional ova were laid. 

Dissection approximately two weeks later revealed that none of the 
H. columbia x H. cecropia ova contained embryos. However, 97.5% 
(157 of 161) of the ova resulting from the H. cecropia x H. cecropia 
mating hatched. The resulting offspring were reared through as pure 
H. cecropia. 

Attempts to obtain hybrid livestock to study the various Postmating 
Mechanisms were hindered by the effectiveness of the prezygotic 
mechanisms and the limited supply of breeding stock. Fortunately, on 
2 June 1982, by placing a number of adults in one cage, Tom Carr 
was able to obtain one pairing of H. cecropia 6 x H. columbia 2? and 
one pairing of H. columbia é x H. cecropia 2 which produced viable 
ova. 

A high degree of mortality during embryonic development was ob- 
served in the hybrid ova. Only 29.1% (7 of 24) of the ova laid by the 
cross-mated H. columbia female hatched and 17.8% (18 of 101) of the 
ova laid by the cross-mated H. cecropia female hatched. In both cases, 
the females laid far less than a full complement of ova. Yet dissection 


78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of collapsing ova from each lot indicated that all of the ova contained 
embryos. 

The loss of hybrids due to larval reluctance to accept potential food- 
plants was also noted. Hybrid larvae were sleeved-out® on several lo- 
cally preferred parental larval foodplants (Tables 2A & 2B). Wild cher- 
ry (Prunus serotina) was given to larvae from both crosses, since it 
appears to be the best foodplant on which to rear Hyalophora hybrids 
(Sweadner, 1937; Wright, 1971). In addition, the hybrid larvae were 
reared in close proximity and under the same conditions as a control 
group of pure H. cecropia larvae. Based upon observations of the hy- 
brid groups and the control group, it did not appear that external 
factors played a significant role in hybrid larval mortality. 

Table 2A depicts the development of hybrids resulting from the H. 
cecropia 6 x H. columbia 2 cross. Since H. columbia females naturally 
Oviposit only on larch, the seven larvae from this cross were offered 
only larch and wild cherry. The five larvae offered larch died in the 
early instars after very little feeding. The two larvae reared on wild 
cherry grew without apparent difficulty and spun cocoons. 

Table 2B shows the development of hybrids resulting from the H. 
columbia é x H. cecropia ? cross. In addition to wild cherry, the 18 
larvae from this cross were offered black willow (Salix nigra), gray 
dogwood (Cornus racemosa), and silver maple (Acer saccharinum). 
The five larvae offered silver maple all died in the Ist instar without 
any evidence of feeding. The five larvae on gray dogwood fed slowly 
and all of them died by the 4th instar. Some degree of success was had 
with black willow, since three of the five larvae spun cocoons. The 
three larvae offered wild cherry completed their development and 
spun cocoons. 

Inviability of the F, hybrids also became apparent. During the fall 
of 1982 the eight hybrid cocoons resulting from the two crosses (Tables 
2A & 2B) were stored with the caged group in Sheridan, Michigan. By 
mid-summer of 1983 only one hybrid adult had emerged from the 
cocoons. Upon cutting the remaining cocoons open, it was determined 
that five of the hybrids had failed to pupate and died in their cocoons 
as larvae. The other two hybrids had successfully pupated but the 
adults had not been able to escape from their cocoons. 

A hybrid female which emerged on 14 June 1983, was the only 
resulting adult from the two crosses. During the female’s lifespan no 
attempts were made at calling. Later dissection revealed that the fe- 
male did not contain ova. 


°A four foot cylinder of nylon screening is extended over a branch of the living foodplant, the larvae are placed 
inside, and boths ends are tied off. 


VOLUME 39, NUMBER 2 79 


TABLE 2A. Comparative development of H. cecropia 6 x H. columbia 2 hybrids 
reared on different larval foodplants. 


N = 
penal Larval Pupal 
ova Foodplant development development F, adults 
2 Wild cherry both completed 1 pupated 1 male failed to 
Prunus serotina development 1 died in cocoon _ escape from cocoon 
as larva 
5 Larch 4 died in 1st instar — _ 
Larix laricina 1 died in 2nd instar 
DISCUSSION 


The results indicate that the three general categories of isolating 
mechanisms (Precontact, Contact, and Postmating) form two indepen- 
dently operating systems. The prezygotic mechanisms (Precontact and 
Contact) limit interaction between H. cecropia and H. columbia by 
controlling adult activities. By restricting interspecific matings and the 
actual exchange of gametes, the reproductive potential of the individ- 
ual female is protected, and more importantly, the opportunity for 
introgression is minimized. 

In contrast, the loss of H. cecropia x H. columbia hybrids due to 
reduced viability is the result of genetic incompatibility. Although fur- 
ther limiting the possibility of breeding F, hybrid adults, it is doubtful 
that postzygotic (Postmating) barriers evolved as a back-up system for 
the prezygotic isolating mechanisms. 

A step by step analysis of Table 1 identifies individual isolating 
mechanisms but also emphasizes their effectiveness when acting in a 
synergistic manner. As a result of its “front-line”? nature, seasonal iso- 


TABLE 2B. Comparative development of H. columbia 6 x H. cecropia 2 hybrids 
reared on different larval foodplants. 


Number Larval Pupal 
of ova Foodplant development development F, adults 
3 Wild cherry all completed 2 pupated 1 female failed 
Prunus serotina development 1 died in cocoon to escape from 
as larva cocoon 
1 female emerged 
on 14 June 1983 
5 Black willow 1 died in lst instar 3 died in cocoon — 
Salix nigra 1 died in 2nd instar as larvae 
5 Gray dogwood 8 died in Ist instar — -— 


Cornus racemosa 1 died in 3rd instar 
1 died in 4th instar 


5 Silver maple all died in Ist — — 
Acer saccharinum instar 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lation is the single-most effective mechanism in limiting interaction 
between H. cecropia and H. columbia, since interspecific contact is 
avoided and mate discrimination is not required. The early phase of 
the H. cecropia flight period in Montcalm County represents only a 
small part of the total population: 18.8% in 1982 and 6.4% in 1983. 
This flight pattern is similar to previously observed bimodal H. cecro- 
pia populations in Missouri and Illinois (Rau & Rau, 1912, 1914; Marsh, 
1941; Sternburg & Waldbauer, 1969). Since the H. columbia flight 
period overlaps only the early H. cecropia flight phase, the potential 
for interaction between the species is substantially reduced (Figs. 2A, 
2B & 3A, 3B). 

A dramatic change in temperature was recorded at the U.S. Weather 
Service reporting station in Greenville, Michigan (approximately 17 
km from the trap site), in the month of May 1982, and the month of 
May 1983. This change presented an opportunity to examine the va- 
lidity of seasonal isolation as an annually utilized isolating mechanism 
in Montcalm County. The monthly mean temperature for May 1982 
was 17.7°C. However, May 1983 was unseasonably cool with a monthly 
mean temperature of only 11.7°C. As a result of the temperature drop 
in 1983, the arrival of wild Hyalophora males at the trap and the 
emergence of the caged group began two weeks later than in the 
previous year. In spite of this, the positioning of the H. cecropia flight 
period with respect to the H. columbia flight period remained constant 
during both years, the entire pattern shifting two weeks to the right 
during the temperature delayed 1983 season. As a result, any interac- 
tion between the two species and all further discussions involve only 
those H. cecropia of the early flight phase and H. columbia. 

The second facet of seasonal isolation takes a closer look at the po- 
tential for interaction by analyzing male-female availability. Rau and 
Rau (1914) noted that H. cecropia males consistently emerged several 
days earlier than H. cecropia females. Prior to the present study, the 
author had made similar casual observations in both H. cecropia and 
H. columbia. 

This mechanism does not minimize contact between H. columbia 
males and H. cecropia females since both are emerging at the same 
time of the year (Figs. 2C & 3C). However, careful analysis of the 
cage emergences indicates that the potential for the occurrence of the 
H. cecropia 6 x H. columbia 2 cross is greatly reduced by the nature 
of male-female emergence patterns. A gap of four days (1982) and 
seven days (1983) existed between the emergence of the last H. cecro- 
pia male from the early emergence phase and the first H. columbia 
female in the caged group during the two years of the study (Figs. 2C 
& 3C). Such a seemingly insignificant gap is very important in affecting 


VOLUME 389, NUMBER 2 81 


the mating behavior of the short-lived Hyalophora adults. Therefore, 
the potential for contact between H. cecropia males and H. columbia 
females rapidly decreases as each day passes, the H. cecropia males 
dying out while the emergence of H. columbia males is peaking and 
the emergence of H. columbia females is just beginning. 

For those individuals emerging during the brief seasonal overlap, 
the subtle variation in daily flight activity of each species serves as the 
next step in reproductive isolation. Variation in circadian rhythm has 
been well documented as an effective isolating mechanism in Callo- 
samia (Brown, 1972; Ferguson, 1972; Peigler, 1980(81)) and Hemileu- 
ca (Collins & Tuskes, 1979). 

The mating activity of Hyalophora adults appears to be closely con- 
trolled by some type of internal biological clock, H. cecropia activities 
beginning at approximately 0330 h’ while H. columbia activities are 
delayed until approximately 0500 h (Figs. 4 & 5). As a result of be- 
coming active prior to the H. columbia adults, there is a period of 
time in which conspecific H. cecropia matings are almost exclusively 
favored. While unmated H. cecropia females were observed calling 
well after dawn, it must be remembered that such behavior by tied 
females was artificially induced at the second Stanton site. 

Conversely, by the time the H. columbia females begin calling, the 
availability of unmated H. cecropia males may be reduced significantly 
by pairing. It is not clear whether the limited H. cecropia male re- 
sponse after the commencement of H. columbia flight activity is the 
result of mating or circadian rhythm terminating flight activity. 

Seasonal and diurnal differences between H. cecropia and H. colum- 
bia mating behavior are very important for maintaining reproductive 
isolation, since no pheromone specificity appears to exist in Hya- 
lophora. The significance of these mechanisms in avoiding interspecific 
contacts can not be overemphasized when considering calling females. 
Interspecific contacts may result in the death of the female or the 
successful exchange of gametes. 

When individuals overcome these Precontact Mechanisms and in- 
terspecific mating is attempted, the incompatibility of genitalia poses 
a further hurdle to successful pairing. Sweadner (1937) noted that less 
than two hours is sufficient to complete fertilization in Hyalophora. 
Yet even in the H. columbia 6 x H. cecropia 2? pairing which lasted 
almost five hours, gametes were not exchanged. While there are minor 
structural genitalic differences between H. cecropia and H. columbia 


7 Exceptions have been noted in the literature (Sweadner, 1937; Ferge, 1983). In addition, recent observations by 
Quimby Hess, Les Kohalmi, and Norm Tremblay (pers. comm.) indicate that H. cecropia flight activity routinely occurs 
between 2100-0200 h in southcentral Ontario. 


82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Ferguson, 1972), the variation in overall size appears to be the single 
most restrictive factor to successful copulation. 

In addition, the tendency of a cross-mated female to re-mate with 
a male of her own species was noted in the present study and has also 
been observed by other investigators (Weast, 1959; Les Kohalmi, pers. 
comm.). The ability of re-mated females to produce pure offspring 
makes this factor very significant. 

On those occasions when a successful interspecific mating occurs and 
gametes are transferred to the female, the various Postmating Mech- 
anisms further restrict the potential for F, hybrid adults. In contrast to 
the prezygotic mechanisms, which maintain the integrity of each gene 
pool by avoiding interspecific matings, postzygotic mechanisms may 
prevent the development of hybrids but at the expense of the repro- 
ductive potential of the individual female. Yet these barriers also pro- 
tect the gene pools by limiting (albeit not eliminating) the successful 
development of hybrids. While previous knowledge of prezygotic 
mechanisms was limited, a great deal of information about postzygotic 
mechanisms was learned by collectors attempting to obtain hybrid 
Hyalophora adults in the lab. As a result, investigators learned that the 
survival rate of F, hybrids varies greatly from one interspecific pairing 
to the next (Sweadner, 1937; Weast, 1959; Kohalmi & Moens, 1975). 

During the present study embryonic mortality claimed a large per- 
centage of the hybrid ova. Although dissection revealed that fertility 
was nearly complete, eclosion of the ova only reached 29.1% in the H. 
cecropia 6 x H. columbia 2 cross and 17.8% in the H. columbia 6 x 
H. cecropia 2 cross. The variability, not only in eclosion but also in 
fertility, between individual cross-matings has been noted by several 
other investigators (Sweadner, 1937; Tom Carr, Ted Herig & Les Ko- 
halmi, pers. comm.). 

By ovipositing on a locally preferred parental foodplant, a cross- 
mated female inadvertently may be further reducing the chance of 
survival for her hybrid progeny. It was noted by the author and by 
previous investigators (Packard, 1914; Sweadner, 1937; Wright, 1971) 
that wild cherry is an excellent foodplant for rearing hybrid Hya- 
lophora larvae. However, it also became obvious that other natural 
larval foodplants successfully utilized by the parent species would not 
effectively support H. cecropia x H. columbia larvae (Tables 2A & 
2B). 

The body of existing literature seems to indicate that the ability of 
H. cecropia x H. columbia hybrids to survive is greatly reduced by 
inviability in all developmental stages (Sweadner, 1937; Weast, 1959). 
The results of the present study would certainly support this conclusion, 
i.e., eclosion of only a small percentage of the fertile ova, a high degree 


VOLUME 39, NUMBER 2 83 


of unsuccessful pupation, and the inability of adults to escape from 
their cocoons. 

Since only one hybrid adult was obtained during the study, no at- 
tempt was made to examine sterility or variation in temporal activity. 
However, the existing literature contains information which helps com- 
plete Table 1. 

Varying degrees of sterility have been reported in F, hybrid Hya- 
lophora adults. While it is generally agreed that F, hybrid males are 
fertile (Sweadner, 1937; Weast, 1959; Collins & Weast, 1961), varied 
results have been obtained with F, hybrid females. Earlier investigators 
(Sweadner, 1937; Freeman, 1940; Weast, 1959; Collins & Weast, 1961) 
all reported F, hybrid females to be totally barren. Recent results (Tom 
Carr & Ted Herig, pers. comm.) indicate that partial fertility is occa- 
sionally obtained in F, hybrid females backcrossed to one of the parent 
species. 

An overview of the results of the study indicates that, at least in 
Montcalm County, Michigan, the occurrence of hybrid Hyalophora 
adults is greatly reduced by a series of isolating mechanisms acting in 
concert. The significance of individual mechanisms may vary from 
year to year and locale to locale. A comparison with other interacting 
Hyalophora populations would be very interesting. 


ACKNOWLEDGMENTS 


Special thanks go out to LaVerne and Evella Petersen of Stanton, Michigan, whose 
maintenance and daily attendance of the bait-trap made the entire project possible. Norm 
Myers of Sheridan, Michigan, Tom Carr of Swanton, Ohio, and Dana Gring of Toledo, 
Ohio, assisted in various aspects of the field work and rearing. M. C. Nielsen of Lansing, 
Michigan, Ted Herig of DeWitt, Michigan, Bruce Wilson of Owosso, Michigan, and 
Charley Chilcote of Cadillac, Michigan, supplied distributional data. The manuscript 
was reviewed and helpful suggestions were offered by Paul Tuskes of Houston, Texas, 
and Michael Collins of Nevada City, California. 


LITERATURE CITED 


BARNES, B. V. & W. H. WAGNER, JR. 1981. Michigan trees; a guide to the trees of 
Michigan and the Great Lakes region. University of Michigan Press, Ann Arbor, 
Michigan. 388 pp. 

BROWN, L. 1972. Mating behavior and life history of the sweetbay silk moth (Callo- 
samia carolina). Science 176:73-75. 

CoLLINs, M. M. 1973. Notes on the taxonomic status of Hyalophora columbia (Satur- 
niidae). J. Lepid. Soc. 27:225-285. 

CoLuins, M. M. & P. M. TuUSKES. 1979. Reproductive isolation in sympatric species of 
dayflying moths (Hemileuca: Saturniidae). Evolution 33:728-733. 

COLLINS, M. M. & R. D. WEAsT. 1961. Wild silk moths of the United States. Collins 
Radio Co., Cedar Rapids, Iowa. 138 pp. 

FERGE, L. A. 1983. Distribution and hybridization of Hyalophora columbia (Lepidop- 
tera: Saturniidae) in Wisconsin. Great Lakes Entomol. 16:67-71. 

FERGUSON, D. C., in R. B. DOMINICK ET AL. 1972. Saturniidae (in part), Bombycoidea, 
The moths of America north of Mexico, fase. 20.2B:155-—275. 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


FREEMAN, T. N. 1940. Notes on the occurrence of Platysamia columbia Sm. in the 
Ottawa region (Lepid., Saturniidae). Can. Entomol. 72:129-130. 

KOHALMI, L. & P. MOENS. 1975. Evidence for the existence of an intergrade population 
between Hyalophora gloveri nokomis and H. columbia in northwestern Ontario 
(Lepidoptera: Saturniidae). Can. Entomol. 107:793-799. 

MaRsH, F. L. 1941. A few life-history details of Samia cecropia within the southwestern 
limits of Chicago. Ecology 22:331-337. 

Mayr, E. 1970. Populations, species, and evolution; an abridgement of animal species 
and evolution. Belknap Press of Harvard University, Cambridge, Massachusetts. 
453 pp. 

MILLER, T. A., J. W. HIGHFILL & W. J. COOPER. 1982. Relationships between pupal 
size and sex in giant silkworm moths (Saturniidae). J. Lepid. Soc. 36:207-216. 

MoorE, SHERMAN. 1955. An annotated list of the moths of Michigan exclusive of 
Tineoidea (Lepidoptera). Miscellaneous Publication No. 88, Museum of Zoology, 
University of Michigan. 87 pp. 

MorTon, E. L. 1895. Hybrid saturniid moths and their larvae. Proc. Entomol. Soc. 
London, pp. 34-35. 

PACKARD, A. S. 1914. Monograph of the bombycine moths of North America, part III. 
Mem. Nat. Acad. Sci., Vol. 12:1-276, Washington, D.C. 

PEIGLER, R. S. 1977. Hybridization of Callosamia (Saturniidae). J. Lepid. Soc. 31: 
23-34. 

1980(81). Demonstration of reproductive isolating mechanisms in Callosamia 
(Saturniidae) by artificial hybridization. J. Res. Lepid. 19:72-81. 

RAu, P. & N. Rav. 1912. Longevity in saturniid moths: An experimental study. J. Exp. 
Biol. 12:179-204. 

1914. Longevity in saturniid moths and its relation to the function of repro- 
duction. Trans. Acad. Sci. St. Louis 23:1-78. 

SMITH, S. I. 1863. Description of a species of Samia, supposed to be new, from Norway, 
Maine. Proc. Boston Soc. Nat. Hist. 9:342-346. 

SOULE, C. G. 1907. Some experiments with hybrids. Psyche 14:116-117. 

STERNBURG, J. G. & G. P. WALDBAUER. 1969. Bimodal emergence of adult cecropia 
moths under natural conditions. Ann. Entomol. Soc. America 62:1422-1429. 

SWEADNER, W. R. 1937. Hybridization and the phylogeny of the genus Platysamia. 
Ann. Carnegie Mus. 25:163-242. 

TiETz, H. M. 1972. An index to the described life histories, early stages and hosts of 
the macrolepidoptera of the continental United States and Canada (in part), Vol. 1. 
Allyn Museum, Sarasota, Florida. 536 pp. 

WEAST, R. D. 1959. Isolation mechanisms in populations of Hyalophora (Saturniidae). 
J. Lepid. Soc. 138:212-216. 

WRIGHT, D. A. 1971. Hybrids among species of Hyalophora. J. Lepid. Soc. 25:66-78. 








Journal of the Lepidopterists’ Society 
89(2), 1985, 85-94 


THE BIOLOGY AND IMMATURE STAGES OF 
SPHINGICAMPA ALBOLINEATA AND S. MONTANA IN 
ARIZONA (SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge #141G, Houston, Texas 77054 


ABSTRACT. The biology and immature stages of Sphingicampa abolineata and 
Sphingicampa montana are described for the first time. The larvae of both species have 
five instars. Development is rapid with only 34-40 days required to progress from egg 
to adult. Evidence indicates that S. albolineata is multiple brooded. The larval host plant 
in Arizona is probably prairie acacia, Acacia angustissima. The larval host of S. montana 
in Arizona remains unknown but based on a selection of native Arizona legumes offered 
to larvae, only sweet acacia, Acacia farnesiana, was found to be acceptable. 


Biological information regarding Sphingicampa albolineata (Grote 
& Robinson) and Sphingicampa montana (Packard) has been lacking 
due in part to the rarity of both species, their restricted distribution 
within the United States, and a past inability to rear the larvae in 
captivity. Ferguson (1971) noted that nothing was known about the 
biology or immature stages of either species and was able to find only 
six U.S. records for both species combined. The purpose of this paper 
is to present new information on the biology, distribution, and imma- 
ture stages of S. albolineata and S. montana. 


Sphingicampa albolineata 


Sphingicampa albolineata is a Mexican species which extends north 
to southern Arizona and Texas. In Arizona the species has a very lim- 
ited distribution and is a resident species in the Huachuca Mts. which 
straddle Cochise and Santa Cruz cos. Specimens have been collected 
most frequently at Washington Camp, Copper Canyon, Miller Canyon, 
Garden Canyon, and occasionally in Ash Canyon. Most Arizona spec- 
imens have been captured between early July and mid August with 
the majority from the first week of August; there is evidence of a 
second generation which flies in mid September. 

Recent records from Texas and northern Mexico indicate two or 
possibly three generations occur per year in those areas. Current Texas 
records include: Black Gap Refuge, Brewster Co., IV-29-82; Browns- 
ville-Southmost, Hidalgo Co., IV-18-82, VI-2-84, X-7-82, and X-20-74. 
Ferguson (1971) also cited a record for Brownsville, XI-10-28. Speci- 
mens from the Gomez-Farias area in Coahuila, Mexico were collected 
X-10-77. A series from Temoris in Chihuahua, Mexico was captured 
between 19 July and 28 August which is similar to the main flight 
period in Arizona. 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Phenotypically, adults from Arizona, western and southern Texas, 
Chihuahua and Coahuila, Mexico appear indistinguishable. There is, 
however, variation in the male genitalia between some of these pop- 
ulations. Texas and Coahuila specimens have a very distinct long thin 
spine on the valve of the male genitalia. Males from central Chihuahua 
have no spine on the valve, while those from Arizona have a short stout 
spine. Fifteen genitalia were examined, and the pattern appeared con- 
sistent. Because Arizona and Texas populations can be separated on 
the basis of the male genitalia, it is possible that their status may 
change. Therefore, it should be noted that all biological observations 
reported here are based on observations of the Arizona population. 

The habitats in which adults have been taken are quite diverse and 
range from thorn forest and oak grassland to mixed forest. Prairie 
acacia, Acacia angustissima (Mill.), is one of the few legumes found 
at all locations where adults have been captured in the Huachuca Mts. 
Acacia angustissima occurs in southern Arizona, Texas and Mexico 
and is a short, multiple-stemmed thornless species with bipinnately 
compound leaves and white flowers. Seeds of angustissima were col- 
lected in the fall and germinated so that potted plants would be avail- 
able if any females were captured. The following year a female was 
collected at the mouth of Copper Canyon and allowed to oviposit in a 
paper bag. 

The ova are green in color and dorsoventrally compressed forming 
a flattened ovoid with a diameter of 2.4 x 1.9 mm. As the embryo 
develops, small gas bubbles appear before the head and body develop. 
At 29°C the ova hatched in 9-11 days. There are five larval instars, 
and development is rapid, requiring only 5-6 weeks to progress from 
egg to adult. 

In addition to the suspected host, larvae were offered various native 
legumes with mixed results. All larvae perished in the first or second 
instar when offered screwbean mesquite, Prosopis pubescens Benth.; 
Jerusalem-thorn, Parkinsonia aculeata L.; paloverde, Cercidium flo- 
ridum Benth.; or sweet acacia, Acacia farnesiana Willd. Larvae reared 
on honeylocust, Gleditsia triacanthos L., had heavy mortality; most 
survivors were stunted and required 3-6 weeks longer to develop com- 
pared to those reared on A. angustissima. Honeylocust is a common 
host plant for Sphingicampa bicolor (Harris) and S. bisecta (Lintner), 
both of which are from the eastern United States, while mesquite is 
the most frequent host of S. heiligbrodti (Harvey) and S. hubbardi 
(Dyar) from the southwest. Larvae offered A. angustissima developed 
to maturity and pupated in 21 to 28 days without mortality. Prairie 
acacia is presumed to be the native host plant of albolineata in southern 
Arizona. Texas prairie acacia, Acacia texensis Torr. & Gray, has been 


VOLUME 39, NUMBER 2 87 


treated as a subspecies of angustissima and was an equally suitable 
host. 

First instar larvae reared on angustissima cling to the underside of 
the petiole and feed at the base of the leaflet. By the end of the second 
day they are large enough to consume a leaflet without moving com- 
pletely off the petiole. During the 3rd through last instar they chew a 
notch on the underside of the petiole and bend the entire leaf back; 
the petiole gives way at the weak point and easily bends into a “v” 
without separating from the plant (Fig. c). In this manner the larva is 
able to consume the distal-most leaflets without having to crawl to the 
tip. Although feeding damage is light during the first four instars, the 
mature larva will consume all of the edible leaves on three to five 
stems. Larvae prefer to feed on leaves of intermediate age. Old leaves 
are avoided and the new growth is not eaten until those of intermediate 
age are consumed. The larva will often leave the plant and wander on 
the ground in search of a new plant prior to consuming the oldest and 
most discolored leaves. 

While on the plant the larva is difficult to find because of its cryptic 
coloration. The silver dorsal and dorsolateral blade-like scoli break up 
the solid green pattern and give the larva the general appearance of 
the thin bipinnately compound leaves on the acacia. Prior to pupation 
the larval coloration changes from a leaf green to dull green, and 
within hours it leaves the plant in search of a pupation site in the soil. 
The pupae reared from ova collected on 31 July eclosed between 16 
and 24 September. They in turn produced another brood which pu- 
pated in mid October of the same year. 

Based on the mid September emergence of reared material and a 
field record from Miller Canyon (IX-14-71) it is probable that at least 
a partial second flight occurs. Considering the rarity of the moth and 
the lack of spring and fall collecting efforts, it is not surprising that 
additional late summer records are lacking. 

In captivity adults emerged from the soil after sunset to 0500 h, with 
a peak between 2200 and 2400 h. Wing expansion is rapid and adults 
are ready to fly within one hour (Fig. e). Individuals that hatch after 
0300 h usually remained quiescent until the next night. In the field 
moths are generally attracted to black lights between 2300 and 0330 
h. Mating occurs after 2200 h, and the pair remain together until the 
following evening. Females that were allowed to oviposit on caged 
potted plants deposited eggs singly on the underside of the leaflets. 
Larval and adult phenotypes within the Arizona population are very 
uniform. The larval description is based on reared material from ova 
deposited by a female captured at the mouth of Copper Canyon, Co- 
chise Co., Arizona. 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. a-h. a, late second instar Sphingicampa albolineata larva feeding on leaflet. 
b, mature fifth instar S. albolineata larva. ec, fifth instar S. albolineata larva on stem 
while consuming leaflets and petiole, note notched petiole. d, mature fifth instar S. 
montana larva, note differences in length and shape of dorsal scoli and size and number 
of small tubercles on lower lateral surface. Adult females: e, S. albolineata; £, S. mon- 
tana, dark form with numerous brownish-black spots on forewings; g, S. montana, light 
form with few or in some instances no brownish-black forewing spots; h, S. montana, 
typical phenotype. 


VOLUME 39, NUMBER 2 89 


Larval Descriptions 


First instar. Head. Diameter 0.9 mm. Light brown with few short brown secondary 
setae. Dark brown line extends from antenna tapering dorsally to vertex of each lobe; 
frontal area dark brown. Body. Ground color green. Length 8.5-9.0 mm, width 2.0 mm. 
Dorsal and dorsolateral meso- and metathoracic scoli (2.0-2.2 mm) light brown with 
short black spines on shaft; tip of each scoli with small brown bulb each with 2 short 
black spines. Dorsal intersegmental area between meso- and metathoracic segments red- 
dish brown. Abdominal dorsal, dorsolateral and lateral scoli green and raised with thin 
black spine extending from each. Mid-dorsal caudal scolus (0.7 mm) light brown with 
numerous short black spines on shaft. Lateral surface with thin dark bluish green line 
above lateral scoli, extending from abdominal (A) segment 1 to A8. Ventral surface and 
true legs green. Prolegs green with dark green shields. . 

Second instar (Fig. a). Head. Diameter 2.7 mm. Color green with yellow medial and 
black distal stripe extending from antennae, tapering dorsally to vertex of each lobe. 
Body. Ground color green. Length 14-15 mm, width 2.5 mm. Dorsal and dorsolateral 
meso- and metathoracic scoli elongated (1.8—2.0 mm), yellow at base with reddish brown 
shafts. Shaft with numerous short black spines; scoli tip with small brown bulb each with 
2 short black spines. Dorsal intersegmental area between meso- and metathoracic scoli 
maroon. Dorsal abdominal scoli light green with light green blade-like projections. Dorsal 
lateral scoli similar to dorsal scoli but half their size. Lateral scoli yellow and reduced in 
size. Sublateral scoli appear as spines on Al, A2, and A7. All abdominal scoli with short 
single black spine. Mid-dorsal caudal scolus (1 mm) green basally with reddish brown 
shaft; numerous short black spines on shaft. Thin black spiracular and light yellow sub- 
spiracular line that encompasses lateral scoli, extends from Al to A8. Ridge of small light 
yellow tubercles form collar on dorsal anterior portion of prothoracic segment. Numerous 
small light yellow tubercles appear on dorsal, lateral, and ventral surfaces, some of which 
form a ring around base of each proleg. Prolegs green. True legs light tan. 

Third instar. Head. Diameter 2.3-2.5 mm. Color green with light yellow stripe ex- 
tending from antennae, tapering dorsally to vertex of each lobe; ocelli at inferior edge 
of line. Body. Ground color green. Length 19-21 mm, width 4.7 mm. Dorsal meso- and 
metathoracic scoli blue, dorsolateral scoli green. Both dorsal and dorsolateral scoli en- 
larged with short black spines on shaft; tips of scoli with small reddish brown bulb, each 
with 2 small black spines. Dorsal intersegmental area between meso- and metathoracic 
segments maroon. Dorsal and dorsolateral abdominal scoli blade-like, light yellow at base 
with silver shaft. Abdominal lateral and sublateral scoli reduced and light yellow. Mid- 
dorsal caudal scolus green with small green spines. Maroon spiracular line extends from 
Al to A8. White line extends from base of dorsolateral metathoracic scoli to become 
subspiracular line extending from Al to A8. Ridge of small light yellow tubercles form 
collar on dorsoanterior portion of prothoracic segment and ridge between meso- and 
metathoracic dorsal scoli. Ridge of yellow tubercles start just dorsal to maroon spiracular 
line and extends over back on posterior and anterior portions of each abdominal segment. 
Light yellow tubercles form ring around base of each proleg, and form 2 poorly organized 
somewhat diagonal lines between prolegs and lateral scoli. True legs and prolegs green. 

Fourth instar. Head. Diameter 3.2-3.4 mm. Color green with light yellow stripe 
extending from antenna, tapering dorsally to vertex of each lobe. Body. Ground color 
green. Length 31-36 mm, width 6 mm. Dorsal meso- and metathoracic scoli turquoise- 
blue with small black spines on shaft. Dorsolateral meso- and metathoracic scoli green 
with small green spines on shaft. Dorsal intersegmental area between meso- and meta- 
thoracic segments green or with trace of maroon. Dorsal and dorsolateral abdominal scoli 
blade-like with yellow base and silver shaft; mesal portion green. Lateral and sublateral 
abdominal scoli reduced to small yellow spines. Mid-dorsal caudal scolus green with small 
white or yellow spines on shaft. Pinkish red spiracular line extends from Al to A8. White 
line extends from base of dorsolateral metathoracic scoli to become subspiracular line 
extending from Al to A8. Ridge of small light yellow tubercles form collar on dorsoan- 
terior portion of prothoracic segment and ridge between meso- and metathoracic dorsal 
scoli. Ridge of yellow tubercles start just dorsal to spiracular line and extend over back 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on posterior and anterior portions of each abdominal segment. Light yellow tubercles 
form ring around base of each proleg and form 2 poorly organized diagonal lines between 
prolegs and subspiracular line. Series of yellow tubercles on lateral thoracic segments just 
dorsal to true legs forming transverse lines on the ventral segmental area of Al and A2. 
Spiracles orange. True and prolegs green. 

Fifth instar (Figs. b, c). Head. Diameter 4.5-5.5 mm. Color bluish green with light 
yellow stripe extending from antennae, tapering dorsally to vertex of each lobe. Antennae 
light yellow. Clypeus green and cream. Body. Ground color green. Length 54-60 mm, 
width 10-12 mm. Dorsal meso- and metathoracic scoli turquoise-blue with yellow tips; 
shaft smooth or slightly knobbed. Dorsolateral meso- and metathoracic scoli yellow with 
silver base; shaft with short white knobs bearing short setae. A line of yellow or silver 
tubercles cross over the mid-dorsal area forming ridge between meso- and metathoracic 
dorsal scoli. Dorsal intersegmental area between meso- and metathoracic segments green. 
Dorsal and dorsolateral abdominal scoli blade-like with silver shaft; mesal portion green. 
Lateral and sublateral abdominal scoli reduced to yellow knobs with small whitish setae 
extending from each. Mid-dorsal caudal scolus greenish yellow with small knobs on shaft. 
Approximate scoli length: thoracic dorsal and dorsolateral, 6-7 mm; mid-dorsal caudal, 
5-6 mm; dorsal abdominal, 3.5 mm; dorsolateral abdominal, 2 mm. All enlarged scoli 
either curved or oriented with their tips in posterior direction. Purplish pink spiracular 
line extends from Al to A8. White or cream colored subspiracular line extends from base 
of dorsolateral metathoracic scoli to A8. Bridge of small light yellow or silver tubercles 
from collar on dorsal anterior portion of prothorax. A more widely spaced ridge of smaller 
silver tubercles starts just dorsal to spiracular line and extends over back on posterior and 
anterior portions of each abdominal segment. Light yellow tubercles form ring around 
base of each true leg and base and tip of each proleg. A series of silver and yellow 
tubercles form 3 ridges on lateral surface: diagonal ridge above base of prolegs; cradle 
under lateral scoli; on lateral thoracic segments just dorsal to true legs. Ventral surface 
of Al, A2, and frequently A7 and A8 with enlarged yellow tubercles forming prominent 
transverse segmental ridge. Anal shield yellow or yellow and silver with silver tubercles. 
True legs and prolegs green. Spiracles orange. 


Sphingicampa montana 


Although present in portions of northern Mexico, Sphingicampa 
montana has an extremely limited distribution in the United States 
and is known from only a few locations in southern Arizona. Ferguson 
(1971) cited five records for this species, all from Pena Blanca Lake, 
with capture dates from 18 July to 8 August. In addition to Pena Blanca 
Lake, this species has been collected at Sycamore Canyon, Nogales, 
and Patagonia in Santa Cruz Co. and Madera Canyon and Box Canyon, 
Pima Co. The flight season extends from late June to mid August with 
a peak between 26 July and 8 August. 

In Arizona, adults are associated with areas dominated by thorn 
forest. The larval host plant in Arizona remains unknown since larvae 
have not been field collected. First instar larvae were offered a wide 
variety of native legumes in an effort to find a suitable host plant. 
Larvae reared on sweet acacia, Acacia farnesiana Willd. developed 
from egg to adult in six weeks with no mortality. The larvae offered 
other native legumes usually died during the first or second instar; 
these plants included: screwbean mesquite, Prosopis pubescens; Jeru- 
salem-thorn, Parkinsonia aculeata; paloverde, Cercidium floridum; 


VOLUME 39, NUMBER 2 91 


prairie acacia, Acacia angustissima; and Mimosa spp. Some larvae 
were successfully reared to maturity on honeylocust, Gleditsia triacan- 
thos. It is interesting to note that the two legumes accepted by mon- 
tana larvae are either totally unacceptable to albolineata larvae or 
result in stunted growth and high larval mortality. Conversely, mon- 
tana larvae could not be reared successfully on prairie acacia, which 
is the larval host plant for albolineata. Steve Prchal (pers. comm.) has 
collected larvae which are believed to be that of montana in Sonora, 
Mexico. The larvae were feeding on Haematoxylon brasalita and a 
large leaved cassia, Cassia emarginata, and were reared to maturity 
on sweet acacia. 

The larvae reared from eggs deposited on 3 August developed rap- 
idly and emerged as adults between 18 and 27 September and subse- 
quently produced another generation which pupated between 14 and 
23 October. The adults reared from larvae collected in Mexico during 
late August by Prchal emerged beginning on 7 September. There are 
no records which indicate a fall flight, thus, the second brood may 
have been an artifact of the rearing conditions. 

The larval feeding habits, female oviposition and mating behavior 
are similar to those described for albolineata. The only major differ- 
ence is that the petiole of sweet acacia is much shorter than that of 
prairie acacia. As a result, mature montana larvae consumed the entire 
leaf without notching the petiole. On longer petiole leaves, such as 
honeylocust, the petiole was notched. so that it could be bent towards 
the larva. Little phenotypic variation was observed among the larvae 
(Fig. d). Of 78 mature larvae only one individual lacked the enlarged 
silver dorsal and dorsolateral scoli on a single segment. Steve Prchal 
indicated that larvae collected in Mexico have little or no silver col- 
oration but when their offspring were reared on Acacia smallii, a small- 
leaved acacia, the majority had silver on all segments. 

The adults are variable. Some have brown wings and brown bodies; 
some have brown wings and yellow bodies; and some have yellow 
wings and yellow bodies. In addition, the brownish black spots of the 
forewing, which are common on most specimens may be totally absent 
(Fig. g) or so dense as to give the marginal area a blackish appearance 
(Fig. f). The specimens which Ferguson (1971) illustrated in color are 
typical of most wild specimens. The colors of reared specimens are 
richer, and in the females the medial forewing area is much lighter in 
coloration than the basal or marginal area of the wing (Fig. h). Newly 
emerged brown specimens are a deep tan, while yellow ones are ac- 
tually a golden yellow. 

The larval description is based on reared material from ova depos- 
ited by a female captured at Pena Blanca Lake, Santa Cruz Co., Ari- 
zona. 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Larval Descriptions 


First instar. Head. Diameter 0.9 mm. Light brown with few short brown secondary 
setae. Dark brown line extends from antenna tapering dorsally to vertex of each lobe. 
Body. Ground color green. Length 8.0-9.0 mm, width 1.8 mm. Dorsal and dorsolateral 
meso- and metathoracic scoli brownish red with short black spines on shaft; tip of each 
scoli with small black bulb each with 2 short black spines. Abdominal dorsal, dorsolateral 
and lateral scoli green and raised with thin black spine extending from each. Mid-dorsal 
caudal scolus brownish red with numerous short black spines on shaft. Caudal scolus 
about 3 times longer than dorsal abdominal scoli. Lateral surface with thin dark bluish 
green line above lateral scoli, extending from abdominal segment 1 to 8. Ventral surface 
green, true legs light brown or green, prolegs brown. 

Second instar. Head. Diameter 1.8 mm. Color green with yellow medial and black 
distal stripe extending from antennae, tapering dorsally to vertex of each lobe. Body. 
Ground color green. Length 12-14.5 mm, width 2.2 mm. Dorsal and dorsolateral meso- 
and metathoracic scoli elongated and brown with numerous short brown spines on shaft. 
Tip of each with small brown bulb each with 2 short black spines. Dorsal abdominal, 
dorsolateral and lateral scoli light yellow with dark brown blade-like projection extending 
from each, lateral scoli reduced in size. Sublateral scoli appear as short spine on Al, A2, 
and A7. Mid-dorsal caudal scolus brown wtih numerous short brown spines on shaft. 
Thin purplish brown spiracular and light yellow subspiracular line, that encompasses 
lateral scoli, extends from Al to A8. Numerous small light yellow tubercles appear on 
the dorsal, lateral and ventral surfaces. Prolegs green with brown shields, true legs light 
brown. 

Third instar. Head. Diameter 2.4—2.6 mm. Color, green with light yellow, black stripe 
extending from antennae, tapering dorsally to vertex of each lobe; ocelli at inferior edge 
of line. Body. Ground color green. Length 17-19 mm, width 4 mm. Dorsal meso- and 
metathoracic scoli brownish red, dorsolateral thoracic scoli yellowish brown. Both dorsal 
and dorsolateral thoracic scoli enlarged with short black spines on shaft; tips of scoli with 
small brown bulb each with 2 small black spines. Dorsal and dorsal abdominal scoli 
blade-like and silver and green. Abdominal lateral and sublateral scoli yellow and re- 
duced. Mid-dorsal caudal scolus reddish brown with short cream colored spines. Red 
subspiracular line and yellow subspiracular lines extend from base of dorsolateral meta- 
thoracic scoli to A8. Ridge of small light yellow tubercles form collar on dorsoanterior 
portion of prothoracic segment. Ridge of yellow tubercles start just dorsal to spiracular 
line and extend over back on posterior and anterior portions of each abdominal segment. 
Light yellow tubercles form ring around base of each proleg, and scattered on lateral 
surface between prolegs and subspiracular line. True and prolegs green. 

Fourth instar. Head. Diameter 3.4-3.6 mm. Color green with light yellow stripe 
extending from antennae, tapering dorsally to vertex of each lobe. Body. Ground color 
green. Length 31-36 mm, width 5.5 mm. Dorsal meso- and metathoracic scoli blue with 
small black spines on shaft. Dorsolateral meso- and metathoracic scoli green to yellow 
with black or white short spines on shaft. Dorsal and dorsolateral abdominal scoli blade- 
like with silver shaft; mesal portion red. Lateral and sublateral abdominal scoli reduced 
to small red spines. Mid-dorsal caudal scolus red or green with small light yellow spines 
on shaft. Thin pink spiracular line extends from Al to A8. Light yellow line extends 
from base of dorsolateral metathoracic scoli to become subspiracular line extending from 
Al to A8. Ridge of small light yellow tubercles form collar on dorsoanterior portion of 
porthoracic segment. Ridge of yellow tubercles start just dorsal to spiracular line and 
extends over back on posterior and anterior portions of each abdominal segment. Yellow 
tubercles form ring around base of each proleg, and true leg, and scattered on lateral 
surface below subspiracular line. A series of yellow and silver tubercles traverse mid- 
dorsal area of meso- and metathoracic segments forming a line that connects base of 
dorsal thoracic scoli, another series forms transverse line on ventral segmental area of Al 
and A2. Anal shield green with yellow, silver, and red tubercles. Spiracles light brown. 
True legs and prolegs green. 

Fifth instar (Fig. d). Head. Diameter 5.3-5.7 mm. Color bluish green with light yellow 


VOLUME 39, NUMBER 2 93 


stripe extending from antennae, tapering dorsally to vertex of each lobe. Antennae light 
yellow. Clypeus green and cream. Body. Ground color green. Length 54 to 60 mm, 
width 10-12 mm. Dorsal and dorsolateral meso- and metathoracic scoli with green base, 
red shaft and yellow tip; shaft with short red knobs bearing short setae. A line of yellow 
tubercles cross over mid-dorsal area forming ridge between meso- and metathoracic 
dorsal scoli. Dorsal intersegmental area between meso- and metathoracic segments green. 
Dorsal and dorsolateral abdominal scoli somewhat triangular shaped with tips curved 
posteriorly; shaft silver with red tip, mesal portion red. Lateral and sublateral scoli 
reduced to red knob with small black setae extending from some. Mid-dorsal caudal 
scolus red with small red or cream colored knobs on shaft. Approximate scoli length: 
thoracic dorsal and dorsolateral, 5 mm; mid-dorsal caudal, 5.5 mm; dorsal abdominal, 
2.2 mm; dorsolateral abdominal, 1.3 mm. All enlarged scoli are curved with their tips 
oriented posteriorly. Red spiracular line extends from Al to A8. White to yellow subspi- 
racular line extends from base of dorsolateral metathoracic scoli to A8. Ridge of small 
light yellow tubercles form collar on dorsoanterior portion of prothoracic segment. A 
more widely spaced ridge of smaller yellow tubercles start just dorsal to spiracular line 
and extend over back on posterior and anterior portions of each abdominal segment. 
Light yellow and red tubercles form ring around base of each true leg and proleg; forms 
semicircle below reduced red lateral scoli and diagonal pattern above base of prolegs. 
Ventral surface of Al, A2 and frequently A7 and A8 with enlarged red and yellow 
tubercles forming prominent transverse segmental ridge. Anal shield green with yellow 
or yellow and silver tubercles. True legs and prolegs green. Spiracles orange-brown. 


DISCUSSION 


Of the three species of Sphingicampa occurring in Arizona only the 
biology and distribution of S. hubbardi was previously known (Com- 
stock, 1947). Although the general shape and appearance of the larvae 
are similar, there are numerous differences between the mature larvae 
of each species. Last instar albolineata larvae have blue dorsal and 
yellowish green dorsolateral meso- and metathoracic scoli. The dorsal 
and dorsolateral abdominal scoli are silver and yellowish green as is 
the mid-dorsal caudal scolus. The tubercles below the spiracular line 
are yellow and silver. Mature montana larvae have red dorsal and 
dorsolateral meso- and metathoracic scoli. The dorsal and dorsolateral 
abdominal scoli are red and silver and the mid-dorsal caudal scolus is 
red. The tubercles below the spiracular line are yellow and red. Finally, 
mature hubbardi larvae have purplish red and yellow dorsal scoli, and 
green and yellow dorsolateral meso- and metathoracic scoli. The dorsal 
and dorsolateral abdominal scoli are silver with a touch of red or pink 
and the mid-dorsal caudal scolus is purple or green. The tubercles 
below the spiracular line are yellow or yellow and red. During the 
early instars the larvae of albolineata have a dark maroon interseg- 
mental patch on the dorsal surface between the meso- and metatho- 
racic segments; montana and hubbardi larvae lack such a patch. Many 
additional differences between albolineata and montana larvae are 
revealed in the larval descriptions. 

The cryptic coloration of albolineata and montana larvae make 
them very difficult to locate while feeding on acacia. The enlarged 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


blade-like dorsal and dorsolateral scoli break up the solid green pattern 
so that the size of the green patches are similar to those of the leaflets, 
and the silver scoli represent the space between leaflets. Larvae of 
montana on acacia are as difficult to located as those of albolineata, 
but individuals reared on honeylocust were easy to locate. This is be- 
cause the leaf on the honeylocust is 4-6 times larger than the acacia 
leaves. It might be expected that populations which feed on large 
leafed hosts would have some of the silver scoli reduced or absent so 
that the pattern would more closely match that of the leaves. 

During the first four instars the dorsal and dorsolateral scoli of all 
three Sphingicampa species orient at 90° to the body, or slant forward 
(Fig. a). In the fifth instar, the dorsal and dorsolateral scoli are curved 
towards the posterior and a series of posterior slanting tubercles form 
a transverse ridge on the ventral surface of abdominal segments | and 
2 and to a lesser degree on A7 and A8. The change in orientation of 
the prominent scoli and development of the ventral ridges are probably 
adaptations to help the larva enter the soil. Posteriorly oriented spines 
might reduce drag while entering the soil and could act to anchor the 
body and prevent it from slipping backwards as it pushes its way 
through the soil. 


ACKNOWLEDGMENTS 


I would like to thank Steve Prchal for sharing his S. montana host plant data based 
on observations in Sonora, Mexico; Jim Tuttle, Steve McElfresh and Tom Carr for sharing 
ova during our Arizona trip; and James Gillaspy, Ed Knudsen, John Hyatt, and Noel 
McFarland for the loan of specimens and/or location data. I also wish to thank Mike 
Collins and Ann McGowan-Tuskes for their suggestions on the manuscript. 


LITERATURE CITED 


ComsTOck, J. A. 1947. Notes on the early stages of Adelocephala heiligbrodti £. hub- 
bardi Dyar. Bull. So. Calif. Acad. Sci. 46:72-77. 

FERGUSON, D. C. 1971. In, The moths of America north of Mexico. Fasc. 20.2a Bom- 
bycoidea (in part). Classey, London, pp. 1-154. 


Journal of the Lepidopterists’ Society 
39(2), 1985, 95-118 


NEVADA BUTTERFLIES: PRELIMINARY CHECKLIST 
AND DISTRIBUTION 


GEORGE T. AUSTIN 


Nevada State Museum and Historical Society, 700 Twin Lakes Drive, 
Las Vegas, Nevada 89107 


ABSTRACT. The distribution by county of the 189 species (over 300 taxa) of but- 
terflies occurring in Nevada is presented along with a list of species incorrectly recorded 
for the state. There are still large areas which are poorly or not collected. 


Nevada continues as one of the remaining unknown areas in our 
knowledge of butterfly distribution in North America. Although a com- 
prehensive work on the state’s butterflies is in preparation, there is 
sufficient demand for a preliminary checklist to justify the following. 
It is hoped this will stimulate those who have any data on Nevada 
butterflies and their biology to forward such for inclusion in the larger 
study. 

Studies of Nevada butterflies are hampered by a paucity of resident 
collectors, a large number of mountain and valley systems and vast 
areas with little or no access. Non-resident collectors usually funnel 
into known and well worked areas, and, although their data are valu- 
able, large areas of the state remain uncollected. Intensive collecting, 
with emphasis on poorly known areas, over the past seven years by 
Nevada State Museum personnel and associates has gone far to clarify 
butterfly distribution within the state. The gaps in knowledge are now 
more narrowly identifiable and will be filled during the next few sea- 
sons. 

There is no all encompassing treatment of Nevada’s butterfly fauna. 
The only state list is an informal recent checklist of species (Harijes, 
1980). Regional works are those for the Carson Range (Herlan, 1962) 
and Clark County (Austin & Austin, 1980). Many records for the east- 
ern, approximately one-third, of the state are mapped by Stanford (in 
Ferris & Brown, 1981). Otherwise, published Nevada records occur 
scattered in various taxonomic revisions, life history and distribution 
studies and in the season summaries of the Lepidopterists’ Society. 

Data for the present paper were obtained from the following sources: 


(1) Collection of the Nevada State Museum (NSM), Carson City. 

(2) Private collections of G. T. Austin, Las Vegas; J. F. Leser, formerly of Las Vegas; C. 
S. Lawson, Las Vegas; C. Crunden, Las Vegas; S. D. Mattoon, Chico, California and 
C. Hageman, Yuba City, California. 

(3) Collection of the Los Angeles County Museum (Clark Co. material only). 

(4) Collection of the Lake Mead National Recreation Area, Boulder City (in part). 

(5) Collection of the Department of Biological Sciences, University of Nevada, Las Ve- 
gas. 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(6) Collection of the Department of Biology, University of Nevada, Reno. 

(7) Ongoing collections for a Great Basin biogeographic study directed by P. Ehrlich, 
Stanford University, Stanford, California. 

(8) Data supplied by various non-resident collectors (see acknowledgments, some of 
whose specimens have been examined; also included are some second-hand data, 
many of which were kindly supplied by R. E. Stanford). 

(9) Data from literature sources (including those in the season summaries of the Lepi- 
dopterists’ Society, most have been subsequently verified through correspondence). 


It is hoped that most important specimens will be examined before 
publication of the larger work. 

Presentation of distributional data for Nevada in a simplified, yet 
meaningful, manner is difficult. Counties, with few exceptions, are 
huge; some reach asinine proportions when trying to consider distri- 
bution. The worst, Nye County, extends nearly half the length and 
breadth of the state from hot Mojave Desert to alpine conditions in the 
Toiyabe Mountains. For present purposes, Nye County is divided into 
two sections at 38° latitude and the northern and southern portions 
herein designated as Nye N and Nye S, respectively. This essentially 
divides the county between the Great Basin and Mojave deserts. Other 
counties (e.g., Washoe, Elko) also create problems but are not subdi- 
vided here. Carson City was previously Ormsby County. Figure 1 
illustrates the counties, and Table 1 indicates the number of taxa re- 
corded in each. 

Nomenclature generally follows Howe (1975; see Ehrlich & Murphy, 
1981) at the generic level and Miller and Brown (1981) at the specific. 
All taxonomic decisions are the author’s, although in some cases they 
were arrived at after consultation with other, more knowledgeable, 
students of the particular taxon. Generally accepted nomenclature is 
presented without comment. In cases where my concepts run counter 
to those in the literature, brief justification is presented. Manuscripts 
in preparation will amplify and further justify these decisions. A ques- 
tion mark indicates that reported specimens were not examined and 
questionable or that the sample was too small for definite subspecific 
determination. 

In an area as large as Nevada, it is expected that there are a number 
of blend zones between populations of different subspecies. This is, in 
fact, the case. In a checklist of this type, however, it is out of place to 
discuss these. The various populations are herein “pigeon-holed” into 
their “‘best fit’? to available names, and a more thorough discussion will 
await forthcoming papers. 


COUNTY RECORDS OF NEVADA BUTTERFLIES 
HESPERIIDAE 


1. Epargyreus clarus huachuca Dixon—Clark. 
2. Polygonus leo arizonensis (Skinner)—Clark, Elko, Lander, Nye N, Nye S. 


VOLUME 39, NUMBER 2 97 


3. 
Aa. 
Ab. 


Im Ht 


Thorybes pylades (Scudder)—Carson City, Clark, Douglas, Washoe. 

Thorybes mexicana nevada Scudder—Carson City, Douglas, Washoe. 

Thorybes mexicana blanca Scott—Lyon, Mineral(?). This taxon was recently de- 
scribed (Scott, 1981). 

Systasea zampa (W. H. Edwards)—Clark. 

Chiomara asychis georgina (Reakirt)—Clark. 

Erynnis icelus (Scudder & Burgess)—Carson City, Elko, Lander, Nye N, Washoe, 
White Pine. 

Erynnis brizo burgessi (Skinner)—Clark, Lincoln. 

Erynnis telemachus Burns—Clark, Lincoln, White Pine. 

Erynnis propertius (Scudder & Burgess)—Carson City, Douglas, Washoe. 
Erynnis meridianus meridianus Bell—Clark, Lincoln. 

Erynnis pacuvius lilius (Dyar)—Carson City, Douglas, Lyon, Washoe. 

Erynnis funeralis (Scudder & Burgess)—Clark, Nye S. 

Erynnis persius (Scudder)—Carson City, Douglas, Elko, Eureka, Humboldt(?), 
Lander, Nye N, Washoe, White Pine. Burns (1964) presented a valid argument 
for not recognizing subspecies at our present state of knowledge. 

Pyrgus ruralis ruralis (Boisduval)—Douglas, Washoe. 

Pyrgus scriptura (Boisduval)—Clark, Lincoln, Nye N, Nye S, White Pine. 


. Pyrgus communis communis (Grote)—Carson City, Churchill, Clark, Douglas, 


Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 


. Pyrgus communis albescens Pl6tz—Carson City, Clark, Esmeralda, Lincoln, Nye 


N, Nye S. This and the preceding are tentatively considered conspecific based 
mainly on the existence of intermediate populations (Tilden, 1965). 

Heliopetes domicella domicella (Erichson)—Clark. 

Heliopetes ericetorum (Boisduval)—Clark, Douglas, Elko, Esmeralda, Eureka, 
Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, White Pine. 
Pholisora catullus (Fabricius)—Carson City, Douglas, Elko, Lander, Lincoln, Lyon, 
Nye N, Storey, Washoe. 


. Pholisora libya libya (Scudder)—Clark, Lincoln, Nye S, White Pine. 
. Pholisora libya lena (W. H. Edwards)—Churchill, Douglas, Elko, Esmeralda, Eu- 


reka, Humboldt, Lander, Lyon, Mineral, Nye N, Nye S, Pershing, Washoe. 
Pholisora alpheus oricus W. H. Edwards—Churchill, Clark, Douglas, Esmeralda, 
Humboldt, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, Washoe. 

Pholisora gracielae MacNeill—Clark. 

Copaeodes aurantiaca (Hewitson)—Clark, Lincoln. 

Hylephila phyleus muertovalle Scott—Carson City, Churchill, Clark, Elko, Lin- 
coln, Nye S. This was recently named by Scott (1981). 


. Pseudocopaeodes eunus nr. wrightii (W. H. Edwards)—Churchill, Lyon, Washoe. 
. Pseudocopaeodes eunus alinea Scott—Nye S. The Amargosa population is distinct 


from others in the state and appears closest to this recently described taxon (Scott, 
1981). 


. Pseudocopaeodes eunus nr. eunus (W. H. Edwards)—Carson City. The Eagle 


Valley population is distinct from others in Nevada and may be worthy of a name. 


. Hesperia uncas lasus (W. H. Edwards)—Elko, Lander, Lincoln, Lyon, Nye N, 


White Pine. 


. Hesperia uncas macswaini MacNeill—Douglas, Esmeralda, Lyon, Mineral, 


Washoe. 


. Hesperia uncas W. H. Edwards ssp.—Elko, Eureka, Lander, Nye N, White Pine. 


This is the large, bright (vs. pale lasus) phenotype of the central Great Basin 
mountains. 


. Hesperia uncas W. H. Edwards ssp.—Mineral. This insect is small and occurs at 


relatively low elevations in extreme western Nevada and adjacent California. 
Hesperia juba (Scudder)—Carson City, Churchill, Clark, Douglas, Elko, Esmeral- 
da, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, 
Storey, Washoe, White Pine. 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


: WHITE PINE 






HUMBOLDT 


MINERAL 





CA=CARSON CITY 
DO=DOUGLAS 
ST =STOREY 


Fic. 1. Map of Nevada showing counties (dashed line shows division between northern 
and southern Nye County as used herein). 


VOLUME 389, NUMBER 2 99 


TABLE 1. Distribution of number of butterfly taxa recorded in each county of Ne- 
vada. 


29a. 


County Number of species Number of taxa 
Carson City 106 114 
Churchill 76 79 
Clark 121 eZ 
Douglas 110 119 
Elko 111 ]LPARE 
Esmeralda 80 87 
Eureka 85 89 
Humboldt 84 96 
Lander 96 106 
Lincoln 110 119 
Lyon 93 100 
Mineral 90 94 
Nye 116 145 

Nye N 103 119 

Nye S 67 76 
Pershing 74 76 
Storey 73 75 
Washoe ANY 130 
White Pine 105 115 
State 189 308 


Hesperia comma harpalus (W. H. Edwards)—Carson City, Churchill, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Storey, Pershing, Washoe, White Pine. Nevada material has been referred to the 
synonymous idaho (W. H. Edwards) and cabelus (W. H. Edwards). 


. Hesperia comma (Linnaeus) spp.—Clark, Nye S. The Spring Range population is 


distinct from any other known population. 

Hesperia pahaska martini MacNeill—Clark, Lincoln. 

Hesperia lindseyi (Holland)—Washoe. 

Hesperia miriamae MacNeill ssp.—Esmeralda. The White Mountains population 
is distinct from those in the Sierra Nevada. 


. Hesperia nevada nevada (Scudder)—Elko. 
. Hesperia nevada (Scudder) ssp.—Carson City, Clark, Douglas, Lyon, Mineral, Sto- 


rey, Washoe. The Sierra Nevada populations are separable from the more eastern 
ones as previously suggested (MacNeill, 1964). 


. Polites sabuleti sabuleti (Boisduval)—Carson City, Churchill, Douglas, Elko, Es- 


meralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, 
Washoe. The name genoa (Plétz), described from Nevada, is synonymous. 


. Polites sabuleti tecumseh (Grinnell)—Carson City, Douglas, Washoe. 
. Polites sabuleti chusca (W. H. Edwards)—Clark, Lincoln, Nye S. 
. Polites sabuleti (Boisduval) ssp.—Eureka, Lander, Nye N, White Pine. These very 


pallid populations are unlike any others in the state. 


. Polites sabuleti (Boisduval) ssp.—Esmeralda. A distinctive high elevation race oc- 


curs in the White Mountains. 


. Polites sabuleti (Boisduval) ssp.—Elko, Lincoln, White Pine. A blackish phenotype 


in eastern Nevada is very distinctive. 
Polites draco (W. H. Edwards)—Clark. 


. Polites sonora sonora (Scudder)—Carson City, Douglas, Esmeralda, Lyon, Mineral, 


Storey, Washoe. 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


36b. Polites sonora utahensis (Skinner)—Humboldt. 

37. Atalopedes campestris campestris (Boisduval)—Clark, Lincoln, Nye S. 

38a. Ochlodes sylvanoides sylvanoides (Boisduval)—Carson City, Churchill, Douglas, 
Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Storey, Washoe, White Pine. 
These populations may eventually be shown to be distinct enough to warrant 
recognition as different from California populations (fide J. F. Emmel). 

38b. Ochlodes sylvanoides bonnevilla Scott—Elko, Eureka, Humboldt, Lander, Persh- 
ing, White Pine. This pallid taxon was described by Scott (1981). 

39. Ochlodes yuma (W. H. Edwards)—Clark, Elko, Esmeralda, Humboldt, Lander, 
Lincoln, Mineral, Nye N, Nye S, Pershing, Washoe. 

40. Euphyes vestris vestris (Boisduval)—Washoe. The name ruricola (Boisduval) ap- 
parently applies to another taxon (fide J. F. Emmel). 

41. Atrytonopsis python (W. H. Edwards)—Clark. 

42. Lerodea eufala (W. H. Edwards)—Clark, Lincoln, Nye S. 

43. Capodes ethlius (Stoll)—Clark. 

44, Agathymus alliae (D. Stallings & Turner) ssp.—Clark, Lincoln. Material represents 
an unnamed eastern Mojave Desert population. 

45a. Megathymus coloradensis maudae D. Stallings, Turner & J. Stallings—Clark, Es- 
meralda, Lincoln, Nye S. These were referred to as navajo Skinner in Austin and 
Austin (1980). 

45b. Megathymus coloradensis browni D. Stallings & Turner—White Pine. 


PAPILIONIDAE 


46. Parnassius clodius baldur W. H. Edwards—Washoe. This was originally reported 
as altaurus Dyer (1967, Lepid. Soc. Season Summary). 

47. Parnassius phoebus sayii W. H. Edwards—Elko. The synonym rubina Wyatt was 
based on Nevada material. 

48. Battus philenor philenor (Linnaeus)—Clark, Lincoln. 

49. Papilio polyxenes coloro W. G. Wright—Clark, Lincoln, Nye S. The use of this 
combination follows Ferris and Emmel (1982); rudkini F. & R. Chermock is thus 


synonymous. 

50. Papilio bairdii W. H. Edwards—Clark, Esmeralda, Lander, Lincoln, Nye N, White 
Pine. 

51. Papilio oregonius oregonius W. H. Edwards—Elko. This may be a bairdii subspe- 
cies. 


52. Papilio zelicaon nitra W. H. Edwards—Carson City, Churchill, Douglas, Elko, 
Eureka, Humboldt, Lander, Lyon, Nye N, Pershing, Storey, Washoe, White Pine. 
Nevada populations are closer to this rather than California zelicaon Lucas (see 
Fisher, 1977). The form “gothica” Remington has been suggested as occurring in 
the state (e.g., Emmel in Howe, 1975). 

58a. Papilio indra indra Reakirt—Carson City, Douglas, Lyon. 

53b. Papilio indra panamintinus J. Emmel—Clark, Lincoln(?), Nye N(?). Placement 
here is tentative pending further study (fide J. F. Emmel). This subspecies was 
recently described (Emmel, 1981). 

58c. Papilio indra nevadensis T. & J. Emmel—Elko, Esmeralda, Lander, Mineral, Nye 
N, Pershing, White Pine. Material from Esmeralda and Mineral counties is inter- 
mediate towards panamintinus. 

53d. Papilio indra Reakirt ssp.—Clark. The populations in the Sheep and certain other 
Clark County mountains are unlike that in the Spring Mountains. 

54a. Papilio rutulus rutulus Lucas—Carson City, Churchill, Douglas, Elko, Esmeralda, 
Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, Storey, 
Washoe, White Pine. The taxon ammoni Behrens, described from Nevada, appears 
unrecognizable as a subspecies. 

54b. Papilio rutulus arizonensis W. H. Edwards—Clark. The Spring Mountains popu- 
lation is different from those in the rest of the state and comes closest to this weakly 
defined race. 

55. Papilio multicaudata W. F. Kirby—Carson City, Churchill, Clark, Douglas, Elko, 


VOLUME 39, NUMBER 2 101 


56. 


BYE TW 


57b. 


58. 


59a. 


59b. 


60. 


61. 


62. 


63. 


64. 


65a. 


65b. 


66a. 


66b. 


67a. 


67b. 
67c. 
68. 


Eureka, Humboldt, Lander, Lincoln, Lyon, Nye N, Pershing, Storey, Washoe, 
White Pine. 

Papilio eurymedon Lucas—Carson City, Douglas, Elko, Humboldt, Storey, Wash- 
oe, White Pine. 


PIERIDAE 


Neophasia menapia menapia (C. & R. Felder)—Carson City, Douglas, Humboldt, 
Lyon, Mineral, Washoe. 

Neophasia menapia (C. & R. Felder) ssp.—Churchill, Clark, Elko, Eureka, Lander, 
Lincoln, Nye N, White Pine. Central and eastern Nevada material differs consis- 
tently from that from the Sierra Nevada and warrants recognition. 

Pieris beckerii W. H. Edwards—Carson City, Clark, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Pieris sisymbrii elivata Barnes & Benjamin—Carson City, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Nye S, Pershing, 
Storey, Washoe. These were called nominate sisymbrii by Edwards (1884) before 
the description of elivata. 

Pieris sisymbrii Boisduval ssp.—Clark, Lincoln, Nye N, Nye S, White Pine. South- 
ern Nevada material is consistently distinguishable from that of the Rocky Moun- 
tains and most of the Great Basin. 

Pieris protodice Boisduval & Leconte—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 

Pieris occidentalis occidentalis Reakirt—Carson City, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, 
Washoe, White Pine. 

Pieris napi pallidissima Barnes & McDunnough—Elko, White Pine. I prefer to 
recognize pallidissima as distinct from Rocky Mountain macdunnoughi Reming- 
ton. 

Pieris rapae rapae (Linnaeus)—Carson City, Churchill, Clark, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Euchloe ausonides ausonides (Lucas)—Carson City, Churchill, Douglas, Elko, Eu- 
reka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, Washoe, White 
Pine. These may be distinct enough from the nominate to require a new name 
(fide J. F. Emmel). 

Euchloe hyantis lotta Beutenmiiller—Carson City, Churchill, Clark, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Euchloe hyantis (W. H. Edwards) ssp.—Carson City, Douglas. This is the Sierra 
Nevada segregate after Opler (1968). 

Anthocharis cethura C. & R. Felder ssp.—Churchill, Esmeralda, Lyon, Mineral, 
Nye S, Pershing, Washoe. The names caliente W. G. Wright and morrisoni W. 
H. Edwards have been attributed to the Nevada fauna (1967 and 1974 Lepid. Soc. 
Season Summaries). The former appears strictly synonymous with nominate ce- 
thura; the latter is probably distinct but does not occur in the state. Typical cethura 
is restricted to southern California; Great Basin material requires a name (fide J. 
F. Emmel). 

Anthocharis cethura nr. pima W. H. Edwards—Clark, Lincoln, Nye S. I consider 
pima as conspecific with cethura on the basis of apparent intermediate populations. 
The Mojave Desert populations are not like southern Arizona pima. 

Anthocharis sara thoosa (Scudder)—Carson City, Churchill, Clark, Douglas, Elko, 
Esmeralda, Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, 
Storey, Washoe, White Pine. 

Anthocharis sara browningi Skinner—Elko, Humboldt. 

Anthocharis sara stella W. H. Edwards—Carson City, Douglas, Washoe. 
Anthocharis lanceolata lanceolata Lucas—Carson City, Douglas, Washoe. 


102 


69. 


70. 


rile 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Colias philodice eriphyle W. H. Edwards—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. This was referred to nominate philodice 
Godart by Austin and Austin (1980). 

Colias eurytheme Boisduval—Carson City, Churchill, Clark, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Colias alexandra edwardsii W. H. Edwards—Carson City, Churchill, Clark, Doug- 
las, Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, 
Nye S, Pershing, Storey, Washoe, White Pine. Some males from most populations 
have orange discal spots on the secondaries and probably represent intergrades 
towards astraea W. H. Edwards (see Ferris, 1973). The name emilia W. H. Ed- 
wards attributed to the state is synonymous. 

Colias cesonia cesonia (Stoll) —Clark, Esmeralda, Lander, Lincoln, Lyon, Mineral, 
Nye N, White Pine. 

Phoebis sennae marcellina (Cramer)—Clark, Lander, Nye N. 

Eurema mexicana (Boisduval)—Clark, Eureka, Nye N. 

Eurema nicippe (Cramer)—Clark, Lincoln, Nye S. 

Nathalis iole Boisduval—Clark, Elko, Esmeralda, Eureka, Lander, Lincoln, Min- 
eral, Nye N, Nye S, White Pine. 


LYCAENIDAE 


. Lycaena arota virginiensis (W. H. Edwards)—Carson City, Churchill, Douglas, 


Elko, Esmeralda, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, 
Washoe, White Pine. 


. Lycaena arota schellbachi (Tilden)—Lincoln, Nye N, White Pine. 
. Lycaena cupreus cupreus (W. H. Edwards)—Douglas, Washoe. 
. Lycaena cupreus artemisia Scott—Elko. The Great Basin phenotype was recently 


named by Scott (198+). 


. Lycaena editha editha (Mead)—Carson City, Douglas, Humboldt, Lander, Storey, 


Washoe. Scott (1979) considered editha synonymous with xanthoides (Boisduva)). 
I believe them to be no less than semispecies. 


. Lycaena editha nevadensis Austin.—Elko, Humboldt. This phenotype is plainly 


distinct from that in western Nevada and was described by Austin (1984). 


. Lycaena rubidus rubidus (Behr)—Humboldt, Washoe. 
. Lycaena rubidus sirius (W. H. Edwards)—Carson City, Churchill, Douglas, Elko, 


Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, Washoe, 
White Pine. Until the recent revision (Johnson & Balogh, 1977), most material 
from west of the Rocky Mountains (including Nevada) was referred to nominate 


rubidus. 


. Lycaena rubidus nr. monachensis K. Johnson & Balogh—Esmeralda, Lyon. 


Lycaena heteronea heteronea Boisduval—-Carson City, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Lycaena dorcas castro (Reakirt)—Clark, Elko. Scott (1978a) argued that western 
members of this group are helloides. Whatever the case, certain Elko County 
populations are phenotypically distinct from helloides. I thus follow Ferris (1977) 
in treating these as a dorcas W. Kirby. 

Lycaena helloides (Boisduval)—Carson City, Churchill, Clark, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 


. Lycaena nivalis nivalis (Boisduval)—Carson City, Douglas, Esmeralda, Humboldt, 


Washoe. The name ianthe (W. H. Edwards), described from Nevada material, is 
synonymous. 

Lycaena nivalis browni dos Passos—Elko, Lander. 

Hypaurotis crysalus crysalus (W. H. Edwards)—Lincoln. 

Habrodais grunus grunus (Boisduval)—Douglas, Washoe. 


VOLUME 39, NUMBER 2 103 


OT. 


88. 


89a. 


89b. 


90a. 


90b. 
Ola. 


91b. 


92a. 


92b. 


92c. 


93. 


94a. 


94b. 


94c. 


94d. 


95. 
96. 


OF: 


98. 
99. 


Atlides halesus estesi Clench—Clark, Douglas, Lincoln, Nye S. This seems to be 
the currently accepted name for western populations rather than corcorani dos 
Passos. 

Harkenclenus titus immaculosus (W. P. Comstock)—Elko, Eureka, Humboldt, 
Lander, Lincoln, Nye N, Pershing, Washoe, White Pine. Western Nevada popu- 
lations may be distinct enough to warrant recognition. 

Satyrium behrii behrii (W. H. Edwards)—Carson City, Churchill, Clark, Douglas, 
Esmeralda, Lander, Lyon, Mineral, Nye N, Nye S, Pershing, Storey, Washoe. 
Satyrium behrii crossi (Field)—Elko, Eureka, Nye N, Lincoln, White Pine. Most 
eastern Nevada populations are darker and more heavily marked beneath and 
larger than Sierran material. These seem to represent crossi. Most specimens from 
Nye and Lander counties are large but pale beneath. 

Satyrium fuliginosum fuliginosum (W. H. Edwards)—Carson City, Douglas, Lyon, 
Mineral, Washoe. 

Satyrium fuliginosum semiluna Klots—Elko, Eureka, Humboldt. 

Satyrium californica californica (W. H. Edwards)—Elko, Eureka, Humboldt, Lan- 
der, Lincoln, Nye N, Pershing, Washoe, White Pine. This is the dark, non-Sierran 
phenotype which occurs over much of the state. It is similar to material from west 
of the Sierra Nevada and is thus included, for now, in that concept. 

Satyrium californica cygnus (W. H. Edwards)—Carson City, Douglas, Esmeralda, 
Lyon, Mineral, Storey, Washoe. Sierran material is sufficiently different from that 
in the rest of Nevada that the name cygnus is raised from synonymy. 

Satyrium sylvinus sylvinus (Boisduval)—Carson City, Churchill, Douglas, Esme- 
ralda, Humboldt, Lyon, Pershing, Storey, Washoe. The subspecific assignments for 
this species are tentative at best until a thorough revisional study is undertaken. 
There appears to be considerable blending between phenotypes. 

Satyrium sylvinus putnami (Hy. Edwards)—Elko, Lander, Lincoln, Nye N, White 
Pine. 

Satyrium sylvinus (Boisduval) ssp.—Carson City, Churchill, Douglas, Elko, Eureka, 
Humboldt, Lander, Lyon, Mineral, Pershing, Storey, Washoe. This large, pale 
phenotype occurs in, mostly, the river valleys. 

Satyrium tetra (W. H. Edwards)—Carson City, Douglas, Storey, Washoe. The 
name adenostomatis (Hy. Edwards) is apparently synonymous. 

Satyrium saepium saepium (Boisduval)—Carson City, Douglas, Storey, Washoe. 
The name fulvescens (Hy. Edwards) described from Lake Tahoe appears synon- 
ymous. At least four phenotypes of saepium occur in the state. Pending a review 
of the taxon, the listing herein represents the best fit to available names. 
Satyrium saepium provo (Watson & W. P. Comstock)—White Pine. Eastern Great 
Basin and Rocky Mountain populations are separable from those of the Sierras; 
this name then refers to the former. 

Satyrium saepium nr. okanagana (McDunnough)—Elko, Humboldt. These are 
very dark with high contrast beneath. 

Satyrium saepium (Boisduval) spp.—Lincoln. This is a distinctive insect (also known 
from Washington County, Utah) with low contrast beneath and broad white edg- 
ings on the submarginal markings. 

Ministrymon leda (W. H. Edwards)—Clark, Elko, Lincoln, Nye N, Nye S. 
Callophrys dumetorum dumetorum (Boisduval)—Douglas. The name dumetorum 
may actually refer to what is now called viridis (W. H. Edwards) whereby the 
name perplexa Barnes & Benjamin would apply to populations currently referred 
to dumetorum (fide J. F. Emmel). 

Callophrys affinis affinis (W. H. Edwards)—Elko, Eureka, Humboldt, Lander, 
Lincoln, Nye N, White Pine. This taxon apparently intergrades with dumetorum 
further north (Scott & Justice, 1981); in fact the entire green Callophrys complex 
may represent a superspecies (fide J. F. Emmel). 

Callophrys comstocki Henne—Clark, Esmeralda, Lincoln, Mineral, Nye N, Nye S. 
Callophrys lemberti Tilden—Carson City, Churchill, Douglas, Elko, Humboldt, 
Lander, Lyon, Mineral, Pershing, Storey, Washoe. 


104 


100. 


101. 
102a. 


102b. 


108a. 


108b. 


104. 


105. 


106a. 


106b. 
107. 


108. 


109. 
110. 


BE 


Za. 


112b. 


113a. 


113b. 


113c. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Callophrys spinetorum spinetorum (Hewitson)—Carson City, Churchill, Clark, 
Douglas, Elko, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, White Pine. All 
Nevada material is of the nominate race and not like the Rocky Mountain ninus 
(W. H. Edwards) (see Clench, 1981). 

Callophrys nelsoni nelsoni (Boisduval)—Carson City, Douglas, Washoe. 
Callophrys siva rhodope (Godman & Salvin)—Clark, Lincoln, Nye S. Green pop- 
ulations from Nevada are much closer to this recently revived taxon (Clench, 1981) 
than to Colorado examples of nominate siva (W. H. Edwards) to which southern 
Nevada material has been heretofore referred (Austin & Austin, 1980). Clench 
(1981) referred one Clark County specimen to nominate siva; the population, 
however, is closer to rhodope. 

Callophrys siva chalcosiva Clench—Carson City, Churchill, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. This is the widespread, brown, Great Basin phenotype 
recently described (Clench, 1981). 

Callophrys augustus iroides (Boisduval)—Carson City, Douglas, Elko, Eureka, 
Humboldt, Pershing, Storey, Washoe. 

Callophrys augustus(?) (W. Kirby) ssp.—White Pine. A short series from the Snake 
Range either represents an undescribed augustus or Callophrys mossii (Hy. Ed- 
wards). It is tentatively placed in augustus but it somewhat resembles Sierran 
mossii windi (Clench). The latter taxon is considered separable from fotis based 
on pattern and, especially, biological differences. 

Callophrys fotis (Strecker)—Clark, Esmeralda, Lincoln, Nye N, Nye S, White Pine. 
Callophrys eryphon eryphon (Boisduval)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Storey, Washoe, 
White Pine. 

Strymon melinus pudica (Hy. Edwards)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Washoe, 
White Pine. 

Strymon melinus setonia McDunnough—Humboldt. 

Brephidium exilis (Boisduval)—Carson City, Churchill, Clark, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Leptotes marina (Reakirt)—Carson City, Churchill, Clark, Douglas, Elko, Esme- 
ralda, Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, Storey, 
Washoe, White Pine. 

Hemiargus ceraunus gyas (W. H. Edwards)—Clark, Lincoln, Nye N, Nye S, White 
Pine. 

Hemiargus isola alce (W. H. Edwards)—Carson City, Clark, Elko, Esmeralda, 
Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, White Pine. 

Everes amyntula amyntula (Boisduval)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, 
Storey, Washoe, White Pine. 

Celastrina ladon echo (W. H. Edwards)—Carson City, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lyon, Mineral, Nye N, Pershing, Storey, Washoe. 
This and the next have long been specifically called argiolus (Linnaeus). It has 
been argued that ladon (Cramer) is probably correct for North American material 
(Clench & Miller, 1980). 

Celastrina ladon cinerea (W. H. Edwards)—Churchill, Clark, Lander, Lincoln, 
Nye N, Nye S, White Pine. 

Euphilotes battoides glaucon (W. H. Edwards)—Carson City, Douglas, Elko, 
Humboldt, Lyon, Mineral, Pershing, Storey, Washoe. The use of Euphilotes follows 
Mattoni (1977). 

Euphilotes battoides baueri (Shields)—Carson City, Churchill, Clark, Douglas, Es- 
meralda, Humboldt, Lander, Lincoln, Nye N, Nye S. 

Euphilotes battoides intermedia (Barnes & McDunnough)—Carson City, Douglas, 
Washoe. 


VOLUME 89, NUMBER 2 105 


118d. Euphilotes battoides martini (Mattoni)—Clark, Lincoln, Nye S. 

118e. Euphilotes battoides nr. bernardino (Barnes & McDunnough)—Esmeralda, Eu- 
reka, Mineral, Nye N, White Pine. 

113f. Euphilotes battoides (Behr) ssp.—Clark. This population is like the fall flying 
phenotype of the eastern Mojave Desert of California (Emmel & Emmel, 1973; 
Shields, 1975, 1977). It has been referred to as near ellisi (Shields). 

113g. Euphilotes battoides (Behr) ssp.—White Pine. The Baking Powder Flat population 
is distinctive (see Shields, 1975). 

118h. Euphilotes battoides (Behr) ssp.—Churchill, Lander, Lyon, Mineral. This entity 
flies in June and associates with Eriogonum heermannii. It has been variously 
referred to as near ellisi and near bernardino. 

113i. Euphilotes battoides (Behr) ssp.—Nye S. A distinctive insect flying in July in the 
Grapevine Mountains (fide J. F. Emmel). 

114a. Euphilotes enoptes ancilla (Barnes & McDunnough)—Churchill, Elko, Eureka, 
Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, Washoe. 

114b. Euphilotes enoptes enoptes (Boisduval)—Carson City, Douglas, Lyon, Storey, 
Washoe. 

114c. Euphilotes enoptes dammersi (J. A. Comstock & Henne)—Clark, Lincoln. 

114d. Euphilotes enoptes (Boisduval) ssp.—Clark, Nye S. This dark and broad-margined 
phenotype in the Spring Mountains is undescribed. 

114e. Euphilotes enoptes (Boisduval) ssp.—Esmeralda, Nye N. This phenotype, some- 
what intermediate between ancilla and the Spring Mountains population, occurs 
in two areas, one in the White Mountains and the other in the Quinn Canyon 
Range. 

115a. Euphilotes mojave mojave (Watson and W. P. Comstock)—Clark. I follow Mattoni 
(1977) in considering mojave as a distinct species. 

115b. Euphilotes mojave langstoni (Shields)—Esmeralda, Mineral. 

116a. Euphilotes rita pallescens (Tilden & Downey)—Churchill, Elko, Esmeralda, Nye 
N. This taxon is considered subspecies of rita (Barnes & McDunnough) after Mat- 
toni (1977). The Sand Mountain population in Churchill County may be distinct. 

116b. Euphilotes rita elvirae (Mattoni)—Carson City, Lyon, Mineral, Washoe. These 
populations were considered within the variation of pallescens by Shields (1977). 
I believe them closer to elvirae than to pallescens. 

116c. Euphilotes rita mattonii (Shields) —Elko. 

116d. Euphilotes rita emmeli (Shields) —Lincoln. 

117. Euphilotes spaldingi spaldingi (Barnes & McDunnough)—Lincoln, White Pine. I 
follow Mattoni (1977) in retaining this as a separate species. 

118. Philotiella speciosa speciosa (Hy. Edwards)—Churchill, Clark, Esmeralda, Lyon, 
Mineral, Nye N, Nye S, Pershing. The use of this genus follows Mattoni (1977). 

119a. Glaucopsyche piasus piasus (Boisduval)—Carson City, Douglas, Washoe. This species 
is undoubtedly a Glaucopsyche as pointed out by Brown (1971). 

119b. Glaucopsyche piasus nevada F. M. Brown—Churchill, Elko, Esmeralda, Eureka, 
Lander, Lincoln, Lyon, Mineral, Nye N, White Pine. These were called daunia 
(W. H. Edwards) before nevada was described. 

119c. Glaucopsyche piasus toxeuma F. M. Brown—Douglas, Humboldt, Pershing, Washoe. 

120a. Glaucopsyche lygdamus oro (Scudder)—Churchill, Clark, Elko, Esmeralda, Eu- 
reka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, White 
Pine. 

120b. Glaucopsyche lygdamus columbia (Skinner)—Carson City, Douglas, Humboldt, 
Lyon, Mineral, Storey, Washoe. The name orcus (W. H. Edwards) may have been 
applied to Nevada material (see Brown, 1970b). 

120c. Glaucopsyche lygdamus (Doubleday) ssp.—Clark, Lincoln, Nye S. This is the large- 
spotted, desert phenotype with an Astragalus host. 

121. Plebejus idas anna (W. H. Edwards)—Carson City, Douglas, Washoe. The name 
argyrognomon (Berstrasser) is considered synonymous (see News Lepid. Soc., 1983: 
66). 

122a. Plebejus melissa melissa (W. H. Edwards)—Carson City, Churchill, Douglas, Elko, 


106 


122b. 
122c. 


123a. 
123b. 


123c. 


124a. 


124b. 


124c. 


125a. 


125b. 
125e. 


126a. 
126b. 


126c. 


Wilas 


127b. 
128. 


129. 
130. 
13la. 


131b. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Plebejus melissa fridayi F. H. Chermock—Carson City, Douglas, Esmeralda, Min- 
eral, Washoe. 

Plebejus melissa (W. H. Edwards) ssp.—Clark, Lincoln. This is the distinctive small 
phenotype of, at least, the Colorado River drainage. 

Plebejus saepiolus gertschi dos Passos—Elko, White Pine. 

Plebejus saepiolus saepiolus (Boisduval)—Carson City, Douglas, Elko, Esmeralda, 
Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Storey, Washoe, White Pine. 
Plebejus saepiolus (Boisduval) ssp.—Esmeralda. This is an unnamed high elevation 
population in the White Mountains. 

Plebejus icarioides fulla (W. H. Edwards)—Carson City, Churchill, Clark, Doug- 
las, Elko, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. This name is the earliest (fide J. F. Emmel) to refer 
to populations of Great Basin influence with nearly immaculate ventral hindwings 
(see Downey in Brown, 1970b). These have heretofore been called ardea (W. H. 
Edwards). 

Plebejus icarioides (Boisduval) ssp.—Clark, Nye S. The Spring Mountains popu- 
lation is distinctive. This was referred to as evius (Boisduval) by Austin and Austin 
(1980). 

Plebejus icarioides (Boisduval) ssp.—Esmeralda, Mineral. Certain populations in 
these two counties do not belong in any described taxon. 

Plebejus shasta minnehaha (Scudder)—Churchill, Douglas, Elko, Esmeralda, Eu- 
reka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, White Pine. 
Emmel and Shields (1978) suggested that minnehaha was a “catch all” name for 
several distinctive populations. 

Plebejus shasta shasta (W. H. Edwards)—Carson City, Washoe. 

Plebejus shasta charlestonensis Austin—Clark. This was recently described by 
Austin (1980). 

Plebejus acmon lutzi dos Passos—Elko, Humboldt, Pershing, White Pine. 
Plebejus acmon texanus Goodpasture—Clark, Esmeralda, Lander, Lincoln, Nye 
N, Nye S, White Pine. 

Plebejus acmon acmon (Westwood & Hewitson)—Carson City, Churchill, Clark, 
Douglas, Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe. 

Plebejus lupini lupini (Boisduval)—Carson City, Churchill, Douglas, Elko, Esme- 
ralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, Storey, Wash- 
oe, White Pine. 

Plebejus lupini nr. monticola (Clemence)—Clark, Nye S. 

Plebejus franklinii podarce (C. & R. Felder)—Carson City, Douglas, Storey, Wash- 
oe. There is no general concensus as to the proper specific name for this insect (see 
Ferris & Brown, 1981; Miller & Brown, 1981). Previously, both glandon (de Prun- 
ner) and aquilo (Boisduval) have been used. 


RIODINIDAE 


Calephelis nemesis californica McAlpine—Clark. 

Calephelis wrighti Holland—Clark. 

Apodemia mormo mormo (C. & R. Felder)—Carson City, Churchill, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. There are, at least, three different mormo in Nevada. 
With the restriction of the type locality to near Pyramid Lake, Washoe County 
(Miller & Brown, 1981), the small, dark, univoltine (late summer), northern Nevada 
phenotype belongs here. 

Apodemia mormo nr. deserti Barnes & McDunnough—Clark, Esmeralda, Lincoln, 
Nye S. This is the small, pale, multivoltine (or at least vernal), desert associated 
phenotype. The insistence by Opler and Powell (1961) that it does not fall into 


VOLUME 39, NUMBER 2 107 


13le. 


132. 


133. 


134. 


135. 
136a. 


136b. 


137a. 


137b. 


138. 


139a. 
139b. 
139c. 


139d. 


139e. 


140a. 
140b. 
14la. 


141b. 
14lc. 


141d. 


142a. 
142b. 


148a. 


148b. 


their conception of deserti prevents me from definately placing it there as have 
others. I, however, can see no consistent differences. 

Apodemia mormo (C. & R. Felder) ssp.—Clark, Lincoln. This is a large, dark, fall 
univoltine which occurs at moderate elevations. 

Apodemia palmerii palmerii (W. H. Edwards)—Clark, Lincoln, Nye S. The name 
marginalis (Skinner) is synonymous, but this phenotype differs from that further 
east. 


LIBYTHEIDAE 
Libytheana bachmanii larvata (Strecker)—Clark, Lincoln. 


HELICONIIDAE 
Agraulis vanillae incarnata (Riley)—Clark. 


NYMPHALIDAE 


Euptoieta claudia (Cramer)—Churchill, Clark, Lincoln, Nye N, White Pine. 
Speyeria cybele leto (Behr)—Carson City, Douglas, Lyon, Washoe. 

Speyeria leto letona dos Passos & Grey—White Pine. 

Speyeria nokomis nokomis (W. H. Edwards)—Elko, White Pine. The Ruby Valley 
population is somewhat intermediate towards apacheana. Ferris and Fisher (1971) 
discussed the blending of nokomis and apacheana across Utah. 

Speyeria nokomis apacheana (Skinner)—Carson City, Douglas, Lander, Lincoln, 
Lyon, Mineral, Nye N, Washoe, White Pine. 

Speyeria coronis snyderi (Skinner)—Carson City, Churchill, Douglas, Elko, Eu- 
reka, Humboldt, Lander, Lyon, Nye N, Pershing, Storey, Washoe, White Pine. 
Nevada specimens were referred to nominate coronis (Behr) by Edwards (1897) 
before snyderi was described. Material from the Sierra Nevada and associated 
ranges is smaller with a browner disc and may be more closely associated with 
simaetha dos Passos & Grey. 

Speyeria zerene zerene (Boisduval)—Carson City, Douglas, Washoe. 

Speyeria zerene malcolmi (J. A. Comstock)—Douglas, Mineral, Storey. 

Speyeria zerene carolae (dos Passos & Grey)—Clark. 

Speyeria zerene platina (Skinner)—Elko, Nye N, White Pine. 

Speyeria zerene gunderi (J. A. Comstock)—Churchill, Elko, Eureka, Humboldt, 
Lander, Nye N, Pershing, Washoe, White Pine. The types of gunderi are of an 
undoubted zerene. The taxon cynna dos Passos & Grey is considered a synonym 
(see Grey, 1975). 

Speyeria callippe nevadensis (W. H. Edwards)—Carson City, Douglas, Elko, Hum- 
boldt, Lyon, Mineral, Storey, Washoe. 

Speyeria callippe harmonia dos Passos & Grey—Churchill, Elko, Eureka, Lander, 
Lincoln, Nye N, Pershing, White Pine. 

Speyeria egleis egleis (Behr)—Carson City, Douglas, Mineral, Washoe. 

Speyeria egleis linda (dos Passos & Grey)—Elko, Humboldt(?). 

Speyeria egleis utahensis (Skinner)—Elko, Eureka, White Pine. 

Speyeria egleis toiyabe Howe—Lander, Nye N. 

Speyeria atlantis greyi (Moeck)—Elko. 

Speyeria atlantis elko Austin.—Elko. Populations of the dodgei (Gunder) cline 
occur in the northern portion of the county. These have been variously referred 
to as near irene (Boisduval) or near dodgei. It was described by Austin (1983). 
Speyeria mormonia mormonia (Boisduval)—Carson City, Douglas, Washoe. The 
restriction of the type locality of mormonia to western Nevada (Miller & Brown, 
1981) seems reasonable. This relegates arge (Strecker), the name previously applied 
to Nevada material, to synonymy. The synonym montivaga (Behr) has also been 
applied to Nevada material (Holland, 1931). 

Speyeria mormonia artonis (W. H. Edwards)—Elko, White Pine. 


108 


144. 


145a. 


145b. 


146. 
147. 


148a. 


148b. 


149. 


150. 
151. 
152. 
153. 


154a. 
154b. 
leaar 
155b. 
155c. 


155d. 


156. 
157. 


158. 


159a. 


159b. 
159c. 
159d. 
159e. 


160a. 
160b. 


16la. 
161b. 
162a. 


162b. 
162c. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Poladryas arachne arachne (W. H. Edwards)—Clark, Lincoln, Nye N, Nye S, 
White Pine. 

Chlosyne leanira cerrita (W. G. Wright)—Clark, Esmeralda, Lincoln, Nye S. A 
new name might be needed for these populations as cerrita was named from a 
mixed population. 

Chlosyne leanira alma (Strecker)—Carson City, Churchill, Douglas, Humboldt, 
Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, Storey, Washoe, White 
Pine. 

Chlosyne californica (W. G. Wright)—Clark, Lincoln. 

Chlosyne lacinia crocale (W. H. Edwards)—Clark. 

Chlosyne palla (Boisduval) ssp.—Carson City, Douglas, Storey, Washoe. This phe- 
notype has been called whitneyi (Behr). Recent investigations have indicated that 
the insect described as whitneyi is actually what we have known as damoetas 
(Skinner) (fide J. F. Emmel). Thus the Sierra Nevada palla is without a name. 
Chlosyne palla vallismortis (J. W. Johnson)—Clark, Nye S. This may actually be 
closer to acastus or a valid species in itself (fide J. F. Emmel). 

Chlosyne acastus acastus (W. H. Edwards)—Carson City, Churchill, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Chlosyne neumoegeni neumoegeni (Skinner)—Clark, Esmeralda, Lincoln, Nye S. 
Chlosyne hoffmanni hoffmanni (Behr)—Carson City, Washoe. 

Phyciodes texana texana (W. H. Edwards)—Clark. 

Phyciodes phaon (W. H. Edwards)—Clark. 

Phyciodes tharos distincta Bauer—Clark. 

Phyciodes tharos nr.(?) pascoensis (W. G. Wright)—Elko, White Pine. 

Phyciodes pratensis pratensis (Behr)—Elko, Esmeralda, Eureka, Humboldt, Lan- 
der, Lyon, Nye N, Pershing, Washoe. 

Phyciodes pratensis montana (Behr)—Carson City, Douglas, Lyon, Mineral, 
Washoe. 

Phyciodes pratensis camillus W. H. Edwards—Elko, Eureka, Lincoln, Nye N, 
White Pine. 

Phyciodes pratensis (Behr) ssp.—Elko, Eureka, Humboldt, Lander. This is a very 
pallid phenotype from the Humboldt River Valley. 

Phyciodes orseis herlani Bauer—Carson City, Douglas, Washoe. 

Phyciodes pallida barnesi Skinner—Clark, Elko, Eureka, Lincoln, Nye N, White 
Pine. 

Phyciodes mylitta mylitta (W. H. Edwards)—Carson City, Churchill, Clark, Doug- 
las, Elko, Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Euphydryas anicia alena Barnes & Benjamin—Clark, Lincoln. The taxa anicia 
(Doubleday & Hewitson) and colon (W. H. Edwards) were synonymized with 
chalcedona (Doubleday) by Scott (1978b). I treat them as, at least, semispecies. 
Euphydryas anicia macyi Fender & Jewett—Humboldt. 

Euphydryas anicia morandi Gunder—Clark. 

Euphydryas anicia veazieae Fender & Jewett—Humboldt, Washoe. 

Euphydryas anicia wheeleri (Hy. Edwards)—Churchill, Douglas, Elko, Esmeralda, 
Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, White Pine. 
Euphydryas chalcedona kingstonensis T. & J. Emmel—Clark. 

Euphydryas chalcedona macglashanii (Rivers)—Carson City, Douglas, Lyon, Sto- 
rey, Washoe. The name truckeensis Gunder, ascribed to our fauna (Martin & 
Truxal, 1955), is synonymous. 

Euphydryas colon nevadensis Bauer—Elko. 

Euphydryas colon wallacensis Gunder—Washoe. 

Euphydryas editha aurilacus Gunder—Washoe. This population has previously 
been referred to nubigena (Behr). 

Euphydryas editha hutchinsi McDunnough—Elko. 

Euphydryas editha lehmani Gunder—Elko, Eureka, Lander, Lincoln, Nye N, White 
Pine. The name caverna Gunder is based on an aberration from Nevada. 


VOLUME 39, NUMBER 2 109 


162d. 


162e. 


162f. 


163. 


164. 


165. 


166. 


167. 


168. 


169. 


170. 


Lids 


172. 


178a. 


178b. 


178c. 
174a. 
174b. 


174c. 
175: 


176. 


Euphydryas editha monoensis Gunder—Carson City, Douglas, Washoe. 
Euphydryas editha koreti Murphy & Ehrlich—Lander, White Pine. The high 
elevation populations of the Toiyabe, Snake and Schell Creek ranges are distinctive 
and were described by Murphy and Ehrlich (1983). 

Euphydryas editha (Boisduval) ssp.—Washoe. This undescribed phenotype is like 
certain low elevation Modoc County, California material. This may or may not be 
what is referred to as edithana (Strand) in northwestern Nevada (Bauer in Howe, 
1975). 

Polygonia satyrus satyrus (W. H. Edwards)—Carson City, Clark, Douglas, Elko, 
Eureka, Lander, Lincoln, Lyon, Nye N, Washoe, White Pine. 

Polygonia zephyrus (W. H. Edwards)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 

Nymphalis californica californica (Boisduval)—Carson City, Churchill, Clark, 
Douglas, Elko, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Per- 
shing, Storey, Washoe, White Pine. 

Nymphalis antiopa antiopa (Linnaeus)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 

Nymphalis milberti furcillata (Say)—Carson City, Churchill, Clark, Douglas, Elko, 
Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, Storey, Washoe, White Pine. 

Vanessa virginiensis (Drury)—Carson City, Churchill, Clark, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Vanessa cardui (Linnaeus)—Carson City, Churchill, Clark, Douglas, Elko, Esme- 
ralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Per- 
shing, Storey, Washoe, White Pine. 

Vanessa annabella (Field)—Carson City, Churchill, Clark, Douglas, Elko, Esme- 
ralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Per- 
shing, Storey, Washoe, White Pine. The name carye (Htibner) was previously 
misapplied to this taxon. 

Vanessa atalanta rubria (Fruhstorfer)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 

Precis coenia (Hiibner)—Carson City, Churchill, Clark, Douglas, Elko, Esmeralda, 
Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, Pershing, 
Storey, Washoe, White Pine. 

Limenitis archippus nr. archippus (Cramer)—Elko. The Little Salmon River pop- 
ulation is somewhat intermediate towards lahontani but is closest to the nominate. 
Limenitis archippus obsoleta W. H. Edwards—Clark. It appears that hulstii W. 
H. Edwards is insufficiently different to warrant recognition. If valid, the latter 
would apply to Nevada material. 

Limenitis archippus lahontani Herlan—Churchill, Elko, Eureka, Humboldt, Lan- 
der, Lyon, Pershing, Storey, Washoe. 

Limenitis weidemeyerii latifascia E. M. & S. F. Perkins—Churchill, Elko, Eureka, 
Humboldt, Lander, Mineral, Nye N, Pershing, White Pine. 

Limenitis weidemeyerii nevadae (Barnes & Benjamin)—Clark. 

Limenitis weidemeyerii angustifascia (Barnes & Benjamin)—Clark, Lincoln. 
Limenitis lorquini eavesii Hy. Edwards—Carson City, Churchill, Douglas, Es- 
meralda, Humboldt, Lyon, Mineral, Storey, Washoe. This species hybridizes with 
L. weidemeyerii latifascia (not nevadae, contra Miller & Brown, 1981); the hybrid 
was named “‘fridayi” Gunder. These are known from Churchill, Elko, Humboldt 
and Mineral counties. Western Great Basin populations are distinct from nominate 
lorquini and fit the concept of eavesii. In the Pine Forest Range (Humboldt Coun- 
ty), the population is largely “fridayi” and the lorquini appears to be of the sub- 
species burrisoni Maynard. 

Adelpha bredowii eulalia (Doubleday & Hewitson)—Clark, Lincoln, White Pine. 


110 


aire 


178. 


179a. 


179b. 


180a. 


180b. 


181. 


182a. 


182b. 


182c. 


188a. 


183b. 


184a. 


184b. 
185a. 


185b. 
185c. 


186. 
187. 


188. 


189. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


All U.S. records (including Nevada, Holland, 1931) were once included in califor- 
nica (Butler). 


APATURIDAE 


Asterocampa celtis montis (W. H. Edwards)—Clark. I have seen no evidence to 
maintain the multitude of monotypic species in this genus. 


SATYRIDAE 


Cyllopsis pertepida dorothea (Nabokov)—Clark, Lincoln. Use of Cyllopsis follows 
the review by Miller (1974). 

Coenonympha ochracea mono Burdick—Douglas, Lyon, Mineral. This, ampelos 
and california are members of the tullia (Muller) superspecies. 

Coenonympha ochracea W. H. Edwards ssp.—Clark, Elko, Eureka, Lander, Lin- 
coln, Nye N, White Pine. The name brenda W. H. Edwards has often been mis- 
applied (e.g., Brown, 1964), as has the nominate (e.g., Holland, 1931) to the heavily 
ocellated Great Basin phenotype; brenda appears synonymous with california (fide 
R. E. Gray; also dos Passos, 1964). 

Coenonympha ampelos ampelos W. H. Edwards—Carson City, Douglas, Elko, 
Eureka, Humboldt, Lander, Lyon, Nye N, Storey, Washoe. 

Coenonympha ampelos elko W. H. Edwards—Elko, Eureka, Humboldt, Lander, 
White Pine. 

Coenonympha california california Westwood—Clark. 

Cercyonis pegala gabbii (W. H. Edwards)—Carson City, Douglas. These popula- 
tions are often referred to as ariane (Boisduval). The latter refers to certain pop- 
ulations west of the Sierra Nevada. The name gabbii may (or may not) apply to 
this western Great Basin material. 

Cercyonis pegala stephensi (W. G. Wright)—Humboldt, Washoe. The name blan- 
ca T. Emmel & Mattoon is a synonym. 

Cercyonis pegala (Fabricius) ssp.—Elko, Eureka, Humboldt, Lander, Lyon, Min- 
eral(?), Pershing, White Pine. The central Great Basin populations are a distinct 
entity. The populations in Lyon and Mineral counties are similar but may not be 
properly placed here. 

Cercyonis sthenele paulus (W. H. Edwards)—Carson City, Churchill, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Cercyonis sthenele masoni Cross—Clark, Lincoln, Nye N, Nye S. This phenotype, 
closest to masoni, extends into the desert areas of California and may warrant 
taxonomic recognition. 

Cercyonis oetus oetus (Boisduval)—Carson City, Churchill, Douglas, Elko, Es- 
meralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Pershing, 
Storey, Washoe, White Pine. 

Cercyonis oetus pallescens T. & J. Emmel—Lander, Nye N. This taxon, described 
by Emmel and Emmel (1971), was omitted in Miller and Brown (1981). 
Neominois ridingsii stretchii (W. H. Edwards)—Elko, Eureka, Humboldt, Lander, 
Nye N, Washoe, White Pine. 

Neominois ridingsii dionysus Scudder—Elko, Nye N, White Pine. 

Neominois ridingsii (W. H. Edwards) ssp.—Esmeralda, Lyon, Mineral. This is the 
pale, western Great Basin population described by Austin (in press). The occurrence 
of the nominate subspecies in Nevada (Emmel in Howe, 1975) is incorrect. 
Oeneis ivallda (Mead)—Carson City, Washoe. 

Oeneis chryxus chryxus (Doubleday & Hewitson)—Elko, Lincoln, White Pine. 


DANAIDAE 


Danaus plexippus plexippus (Linnaeus)—Carson City, Churchill, Clark, Douglas, 
Elko, Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye 
S, Pershing, Storey, Washoe, White Pine. 

Danaus gilippus strigosus (Bates)—Carson City, Churchill, Clark, Douglas, Elko, 


VOLUME 39, NUMBER 2 EFT 


Esmeralda, Eureka, Humboldt, Lander, Lincoln, Lyon, Mineral, Nye N, Nye S, 
Pershing, White Pine. 


DUBIOUS AND HYPOTHETICAL RECORDS 


A number of taxa have been reported for Nevada which are unlikely 
or represent misdeterminations. Others range nearly to the borders of 
the state and may be expected to occur. These are commented upon 
below. Taxa reported but now considered synonymous with those oc- 
curring in the state are discussed above in the main species accounts. 


Erynnis brizo lacustra (W. G. Wright)—The reported record (Martin & Truxal, 1955) 
undoubtedly refers to burgessi. 

Hesperia comma oregonia (W. H. Edwards)—The type series was supposedly taken 
in Nevada and there are specimens labeled such in Edwards’ collection in the Carnegie 
Museum, Pittsburgh, Pennsylvania (see Brown & Miller, 1977). These probably led to 
the subsequent listing of the taxon for Nevada (Lindsey, 1921; Lindsey et al., 1931). This 
subspecies does not occur in the state. 

Hesperia pawnee Dodge—MacNeill (1964) mentioned seeing an, undoubtedly misla- 
beled, male labeled “Nevada.” 

Hesperia viridis (W. H. Edwards)—A specimen in the Snow Entomological Museum, 
University of Kansas, Lawrence, is labeled “Verdi, Nevada, July, 1903” (MacNeill, 1964; 
Brown & Miller, 1977). It is undoubtedly mislabeled. 

Ochlodes sylvanoides pratincola (Boisduval)—A worn Nevada specimen taken to be 
this taxon was figured by Holland (1931). It probably is nominate sylvanoides. 

Ochlodes agricola (Boisduval)—Specimens listed in the Nevada State Museum catalog 
for Clark and Elko counties are unlocatable. Undoubtedly these represent a misdeter- 
mination of some other taxon. Specimens from the W. H. Edwards collection from 
Nevada (Irwin, 1966) are probably mislabeled. 

Poaenes taxiles (W. H. Edwards)—The taxon has been listed for the state on several 
occasions (e.g., Edwards, 1881; Lindsey, 1921; Lindsey et al., 1931; MacNeill in Howe, 
1975; Pyle, 1981). Three specimens labeled Nevada are in the W. H. Edwards collection 
at the Carnegie Museum, Pittsburgh, Pa. (Brown & Miller, 1980). There are no recent, 
verifiable records for the state but it occurs into western Utah and may be found even- 
tually in one or more of the eastern Nevada counties. 

Paratrytone melane melane (W. H. Edwards)—Nevada, in error, was included as the 
type locality (see Brown & Miller, 1980). The species is unknown in the state. 

Amblyscirtes eos (W. H. Edwards)—The one report for Clark County (1972, Lepid. 
Soc. Season Summary) represents a misdetermination of Pholisora alpheus (fide J. F. 
Lesser). 

Amblyscirtes vialis (W. H. Edwards)—Holland (1931) stated the range as including 
Nevada. I do not know of any records for the state although it occurs not too far away 
in the Sierra Nevada of California (Shapiro et al., 1979). 

Parnassius clodius sol Bryk & Eisner—The type locality was listed as “Nevada’’; this 
is probably more properly the Sierra Nevada somewhere in California. This taxon is not 
known from Nevada. 

Parnassius phoebus behrii W. H. Edwards—Brown (1975b) mentioned a total of seven 
specimens in the W. H. Edwards collection at the Carnegie Museum labeled Nevada. 
There are no recent records and the above may represent mislabeling although the taxon 
occurs close to the Nevada line in the Sierra Nevada. 

Papilio indra fordi J. A. Comstock & Martin—Tyler’s (1975) inclusion of this taxon 
for Nevada is erroneous (probably based on 1963, Lepid. Soc. Season Summary). The 
record undoubtedly refers to the panamintinus-like populations in the Spring Mountains. 

Neophasia menapia tau (Scudder)—The Caron Range population (Herlan, 1962) is 
not of this subspecies but of the nominate. 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Euchloe hyantis hyantis (W. H. Edwards)—The recorded occurrence (1967, Lepid. 
Soc. Season Summary; see also Brown, 1973) refers to lotta. 

Anthocharis sara sara Lucas—This has been incorrectly included in the Carson Range 
list (Herlan, 1962) as the form “reakirtii’”» W. H. Edwards. Two specimens of this taxon 
labeled “Mineral County” are in the Nevada State Museum. They are regarded as mis- 
labeled as they do not, in any way, resemble material from nearby. 

Anthocharis sara inghami Gunder—This and thoosa have been confused leading to 
the erroneous use of the former in the Nevada literature (1969, Lepid. Soc. Season 
Summary). 

Colias occidentalis chrysomelas Hy. Edwards—The report for the Carson Range (Her- 
lan, 1962) is unverified and undoubtedly represents a misdetermination. 

Lycaena xanthoides xanthoides (Boisduval)—This species does not occur in Nevada; 
the Carson Range record (Herlan, 1962) is an undoubted misdetermination. 

Lycaena gorgon (Boisduval)—I do not know the basis for Holland’s (1931) inclusion 
of this species for Nevada. 

Lycaena mariposa mariposa (Reakirt)—This species is reported as occurring in Nevada 
by Opler (in Howe, 1975). I know of no records. 

Satyrium acadica coolinensis (Watson & W. P. Comstock)—The supposed Nevada 
record (Herlan, 1962) is a misidentification of, probably, californica. 

Satyrium sylvinus dryope (W. H. Edwards)—A pair labeled dryope collected in “Ne- 
vada” by Morrison are in Edwards’ collection at the Carnegie Museum (Brown, 1970a). 
These are either mislabeled or represent tailless individuals from a normally tailed Ne- 
vada population. 

Callophrys sheridanii (W. H. Edwards)—Pyle (1981) reported sheridanii for southern 
Nevada. He treated this taxon as specifically distinct from both comstocki and lemberti. 
In this sense, sheridanii is unverified for Nevada. 

Callophrys mossii windi (Clench)—This butterfly is in the Sierra Nevada not far from 
the Nevada line (fide D. L. Bauer). It may occur in association with its Sedum foodplant 
in, especially, the Mt. Rose area of Washoe County. 

Everes comyntas (Godart)—This was reported for Nye County (1969, Lepid. Soc. 
Season Summary). All Nevada Everes seem to be amyntula. 

Glaucopsyche piasus sagittigera (C. & R. Felder)—Brown (1975a) placed a single 
Humboldt County specimen in this taxon. More extensive material from this area shows 
these to be toxeuma. 

Glaucopsyche lygdamus incognitus Tilden—The Nevada occurrences (Martin & Truxal, 
1955; Herlan, 1962) as behrii (W. H. Edwards) probably represent columbia. 

Plebejus icarioides lycea (W. H. Edwards)—The Carson Range report (Herlan, 1962) 
is of fulla. 

Plebejus icarioides icarioides (Boisduval)—Two names associated with this subspecies 
have been ascribed to Nevada. The first is mintha (W. H. Edwards) of which the types 
were originally stated as being from Nevada but later corrected to California (see Brown, 
1970b). The other, fulla (W. H. Edwards), has been synonymized with nominate ica- 
rioides (e.g., dos Passos, 1964; Miller & Brown, 1981). This, however, is the senior syn- 
onym of the widespread Great Basin subspecies previously called ardea. 

Speyeria nokomis nitocris (W. H. Edwards)—This taxon was erroneously reported for 
Nevada (Edwards, 1897; dos Passos & Grey, 1947). 

Speyeria zerene conchyliatus (J. A. Comstock)— Variation among the blending zerene 
populations east of Lake Tahoe produces occasional specimens resembling this subspecies 
(e.g., Herlan, 1962). The variation is best referred to the nominate subspecies. The Sierran 
influence seen in some individuals from northwestern Washoe County is due to introgres- 
sion from conchyliatus. 

Speyeria callippe juba (Boisduval)—This taxon was included for Nevada as inornata 
(W. H. Edwards) by Holland (1931) and dos Passos and Grey (1947), both probably 
following Edwards (1884). I know of no records although it does occur in the Sierra 
Nevada not far to the west (Shapiro et al., 1979). 

Speyeria callippe laura (W. H. Edwards)—The types of this subspecies were reported 


VOLUME 39, NUMBER 2 118 


from Nevada (Edwards, 1879). Nothing like it has turned up in the state since. The name 
probably applies to something further west in California. 

Speyeria atlantis irene (Boisduval)—Moeck (1957) reported irene on Verdi Peak north 
of Lake Tahoe near the Nevada line. It may occur east of here in Washoe County. 

Boloria epithore sierra E. Perkins—The species occurs in the vicinity of South Lake 
Tahoe, Eldorado County, California (fide D. L. Bauer). It may occur in adjacent Douglas 
County. 

Chlosyne leanira wrightii (W. H. Edwards)—Holland (1931) erroneously included 
Nevada in the range of this taxon. 

Chlosyne gabbi (Behr)—Higgins (1960) reported a specimen of this species labeled 
“Nord Nevada.” It is undoubtedly mislabeled. 

Chlosyne whitneyi whitneyi (Behr)—This has been long known as damoetas (Skinner) 
(see comment under palla in main species accounts). The species occurs in the Sierra 
Nevada and Sweetwater Mountains in California. It may also be in adjacent Nevada. 

Dymasia dymas chara (W. H. Edwards)—A specimen in the Allyn Museum of Ento- 
mology, Sarasota, Florida is mislabeled Elko County (fide E. M. Perkins). The correct 
data are Pima County, Arizona, and the above museum has been so notified. 

Phyciodes picta (W. H. Edwards)—This species was erroneously listed for Nevada 
(1964, Lepid. Soc. Season Summary). There are no records for the state. 

Phyciodes pallida pallida (W. H. Edwards)—This taxon was reported from Nevada 
as mylitta mata (Reakirt) (1963, Lepid. Soc. Season Summary). This undoubtedly rep- 
resents barnesi. 

Euphydryas chalcedona olancha (W. G. Wright) and sierra (W. G. Wright)—Holland 
(1931) and Scott (1978b) used these names for the variation in Nevada macglashanii. 
Comstock (1937) referred to central Nevada material (apparently wheeleri) as sierra. 

Polygonia faunus rusticus (W. H. Edwards)—This species has been taken a very short 
distance from the Nevada line near South Lake Tahoe, Eldorado County, California (fide 
D. L. Bauer) and will probably eventually be recorded in Douglas County. 

Polygonia oreas silenus (W. H. Edwards)—Ferris and Brown (1981) showed an un- 
verified record for Elko County. This represents a misdetermined zephyrus (fide R. L. 
Langston). 

Precis evarete (Cramer)—The report for Nevada (Herlan, 1962 as orithya evarete) 
refers to coenia. 

Coenonympha ampelos columbiana McDunnough—Nevada nominate ampelos have 
been erroneously referred to this taxon (Herlan, 1962; 1964, Lepid. Soc. Season Sum- 
mary). 

Cercyonis pegala wheeleri (W. H. Edwards)—The types for the synonymous hoffmani 
(Strecker) were reported as from “Owens Lake, Nevada.” This locality is actually in 
California. 

Cercyonis sthenele silvestris (W. H. Edwards)—The occurrence in Nevada (Herlan, 
1962) refers to paulus. 

Oeneis nevadensis nevadensis (C. & R. Felder)—Martin and Truxal (1955) and Em- 
mel (in Howe, 1975) reported this species for Nevada. I know of no definite records. 

I have been unable to verify a number of possible county records listed in the Harjes 
(1980) checklist. These are listed below. Some are unlikely or are known misdetermi- 
nations and, the county is set in italics, and many are commented upon. Others are 
probable and are listed mostly without comment. 


Hesperia nevada—Esmeralda (misdetermined uncas, NSM). 

Heliopetes ericetorum—Washoe. 

Pyrgus ruralis—Carson City. 

Erynnis propertius—Lyon (misdetermined pacuvius, NSM). 

Thorybes mexicana—White Pine (misdetermined Erynnis telemachus, NSM). 
Papilio bairdii—Washoe (misdetermined zelicaon, NSM). 

Nathalis iole—Douglas. 

Lycaena nivalis—Churchill (misdetermined helloides, NSM). 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Habrodais grunus—Clark (typographical error, should be Douglas). 

Callophrys spinetorum—Storey. 

Callophrys nelsoni—Elko, Lyon, Nye (misdetermined siva, NSM, etc.). 

Callophrys dumetorum—Storey (misdetermined lemberti, Bauer). 

Plebejus idas—Mineral (misdetermined melissa, NSM). 

Libytheana bachmanii—Storey. 

Limenitis weidemeyerii—Carson City, Washoe. 

Limenitis lorquini—White Pine. 

Polygonia satyrus—Storey (misdetermined zephyrus, NSM). 

Chlosyne lacinia—Lander. 

Euphydryas chalcedona—Churchill, Elko, Eureka, Lander, Nye (correct only if anicia 
is considered a Chalcedona, see Scott, 1978b). 

Euphydryas editha—Churchill. 

Danaus gilippus—Washoe. 

Coenonympha california—Humboldt (partially mislabeled California specimen, NSM). 

Neominois ridingsii—Douglas. 
There are an additional five county records that are erroneous or unverified in any 

way: 


Hesperia pahaska—White Pine (Ferris & Brown, 1981). This is possible, but there are 
no verified records to date. 

Chlosyne palla—Elko (Ferris & Brown, 1981). Unverified and probably refers to acastus. 

Limenitis lorquini—Elko, White Pine (Ferris & Brown, 1981). The White Pine record 
is totally erroneous; the Elko record undoubtedly refers to “fridayi’” specimens which 
occasionally turn up. 

Cercyonis oetus—Clark (1974, Lepid. Soc. Season Summary). Undoubtedly this refers 
to sthenele. 


Other Clark County dubia are listed in Austin and Austin (1980). 


DISCUSSION 


To date, 189 species and some 300 total taxa of butterflies are known 
from the state of Nevada (Table 1). In general, counties with a portion 
of the Sierra Nevada in western Nevada, the four counties on the 
eastern border and the huge Nye County show the greatest diversity. 
Some of this is real; some is undoubtedly due to insufficient collecting. 
In addition, there are a number of areas within the state that have 
received little or no study. These include, but are not limited to, the 
following: 


1) the northeastern and southeastern portions of Elko County. 
2) Elko and Humboldt counties between the Independence and Santa 
Rosa ranges. 
3) western Humboldt County west of the Santa Rosa Range. 
4) eastern and western Pershing County. 
) extreme northern Washoe County. 
) much of Churchill County outside the Clan Alpine Range and the 
Fallon area. 
7) Lander County between U.S. 50 and I-80. 
8) northwestern quarter of White Pine County. 
9) northern Nye County except the region from the Toiyabe Range 
to the Monitor Range. 


VOLUME 39, NUMBER 2 ELS 


10) southern Nye County (most of this is due to the presence of the 
Nevada Test Site which is off limits to the average collector; there 
are still fringe areas which can be studied). 

11) western half of Lincoln County. 

12) Mineral County except the Wassuk Range. 

13) Esmeralda County except the White Mountains. 


Some of these areas appear important as blend zones between taxa 
or may represent the distributional limits of others. 

What is as important as filling in the distributional holes in Nevada 
is a more thorough knowledge of the fauna of adjacent regions. A start 
on this is Dornfeld’s (1980) work on Oregon butterflies. The distribu- 
tion and taxonomy of the butterflies of the other bordering states (Ar- 
izona, California, Idaho, Utah) are in various stages of study and up- 
dating. Once completed, we should have a picture of the influence of 
surrounding regions on the butterflies of Nevada specifically and the 
Great Basin in general. 


ACKNOWLEDGMENTS 


Numerous people provided data, suggestions and other help in the preparation of this 
checklist. The group at Stanford University under the direction of P. Ehrlich, including 
D. Murphy, O. Shields and B. Wilcox were most helpful in keeping me abreast of their 
activities and collections. Other records for the state were received from the following: 
D. E. Allen, R. Bailowitz, D. L. Bauer, A. Bean, J. Brock, F. M. Brown, J. M. Burns, C. 
Callaghan, H. Clench (Carnegie Museum records of lycaenids), J. T. Cooper, C. Crunden, 
T. E. Dimock, D. Eff, J. F. Emmel, T. C. Emmel, C. D. Ferris, C. F. Gillette, R. E. 
Gray, L. P. Grey, D. Guiliani, C. Hageman, G. Harjes, C. Henne, P. J. Herlan, H. L. 
King, J. Lane, R. L. Langston, C. S. Lawson, J. F. Leser, A. Ludke, W. W. McGuire, C. 
D. MacNeill, J. Masters, S. O. Mattoon, D. Mullins, J. S. Nordin, F. W. Preston, R. 
Robertson, K. Roever, F. Ryser, J. A. Scott, C. Sekerman, O. E. Sette, O. Shields, D. 
Shillingburg, M. Smith, N. J. Smith, R. E. Stanford, G. B. Straley, W. Swisher, D. Thomas, 
K. B. Tidwell, J. W. Tilden and R. E. Wells. Access to the collections at the Los Angeles 
County Museum was made possible by J. Donahoe, at Lake Mead National Recreation 
Area by D. H. Huntzinger, University of Nevada, Reno by F. Ryser and University of 
Nevada, Las Vegas by C. Murvosh. L. D. Miller sent certain specimens housed at the 
Allyn Museum of Entomology. J. F. Emmel and R. E. Stanford made numerous helpful 
comments and suggestions on a late draft. All of these are gratefully thanked. A number 
of the above also made taxonomic suggestions which were of great help. Last, but not 
least, I thank Pam Church for her patience in typing some of the early drafts of this and 
correcting all my errors and omissions. 


LITERATURE CITED 


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(Lycaenidae). J. Lepid. Soc. 34:20-24. 

1984. A new subspecies of Speyeria atlantis (Edwards) (Nymphalidae) from 

the Great Basin of Nevada. J. Lepid. Soc. 37:244-248. 

1984. A new subspecies of Lycaena editha (Mead) (Lycaenidae) from Nevada. 
J. Res. Lepid. 23:83-88. 

AUSTIN, G. T. & A. T. AUSTIN. 1980. Butterflies of Clark County, Nevada. J. Res. 
Lepid. 19:1-63. 








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VOLUME 39, NUMBER 2 LZ 


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America north of Mexico. Lepid. Soc. Mem., no. 2. 

MokEck, A. H. 1957. Geographic variability in Speyeria. Comments, records and de- 
scription of a new subspecies. Milwaukee Entomol. Soc., Special Paper. 

MurpnHy, D. D. & P. R. EHRLICH. 1983. Biosystematics of the Euphydryas of the 
central Great Basin with the description of a new subspecies. J. Res. Lepid. 22:254- 
261. 

OPLER, P. A. 1968. Studies on Nearctic Euchloe. Part 5. Distribution. J. Res. Lepid. 7: 
65-86. 

OPpLER, P. A. & J. A. POWELL. 1961. Taxonomic and distributional studies on the 
western components of the Apodemia mormo complex (Riodinidae). J. Lepid. Soc. 
15:145-171. 

PyLE, R. M. 1981. The Audubon Society field guide to North American butterflies. A. 
A. Knopf, New York. 

ScoTT, J. A. 1978a. The identity of the Rocky Mountain Lycaena dorcas-helloides 
complex (Lycaenidae). J. Res. Lepid. 17:40-50. 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1978b. A survey of valvae of Euphydryas chalcedona, E. c. colon, and E. c. 

anicia. J. Res. Lepid. 17:245-252. 

1979. Geographic variation in Lycaena xanthoides. J. Res. Lepid. 18:50-59. 

1981. New Papilionoidea and Hesperioidea from North America. Papilio (new 
series), no. 1. 

ScoTT, J. A. & J. A. JUSTICE. 1981. Intergradation between Callophrys dumetorum 
oregonensis and Callophrys dumetorum affinis in northwestern U.S. (Lycaenidae). 
J. Res. Lepid. 20:81-85. 

SHAPIRO, A. M., C. A. PALM & K. L. WcISLO. 1979. The ecology and biogeography of 
the butterflies of the Trinity Alps and Mount Eddy, northern California. J. Res. 
Lepid. 18:69-151. 

SHIELDS, O. 1975. Studies on North American Philotes (Lycaenidae). IV. Taxonomic 
and biological notes, and new subspecies. Bull. Allyn Museum, no. 28. 

1977. Studies of North American Philotes (Lycaenidae). V. Taxonomic and 
biological notes, continued. J. Res. Lepid. 16:1-67. 

TILDEN, J. W. 1965. A note on Pyrgus communis and Pyrgus albescens (Hesperiidae). 
J. Lepid. Soc. 19:91-94. 

TYLER, H. A. 1975. The swallowtail butterflies of North America. Naturegraph, Healds- 
burg, California. 











Journal of the Lepidopterists’ Society 
39(2), 1985, 119-124 


MITOURA MILLERORUM (CLENCH) AND ITS OCCURRENCE 
IN THE UNITED STATES (LYCAENIDAE) 


KURT JOHNSON 


Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024 


ABSTRACT. The male imago is illustrated and the male genitalia described and 
illustrated for the first time. A new Mexican and a thus-far unique southwestern United 
States collection record are documented along with clarifications concerning the original 
description of millerorum, specimens which represent it, and its inter-specific diagnosis. 
A larval specimen collected on Arceuthobium globosum Hawksworth and Wiens (Lo- 
ranthaceae) is discussed relative to its possibly representing millerorum and, if so, the 
importance of any apparent monophagy in defining the species. 


Mitoura millerorum (Clench) was described from a holotype female 
collected in the vicinity of E] Encarnacion, Hidalgo State, Mexico, with 
a male paratype (not personally examined by Clench) designated from 
Palos Colorados, Durango State, Mexico (Clench, 1981). The purposes 
of this brief paper are to: (1) illustrate the male imago and describe 
and illustrate the male genitalia of the hitherto undescribed male of 
millerorum; (2) document further collection records of the species, 
including one in the United States; (3) clarify the comments of Clench 
(loc. cit.) concerning the species and specimens representing it; (4) 
summarize the present knowledge of its biology and (5) thereby en- 
courage lepidopterists in the United States to pursue larger samples of 
this apparently rare hairstreak butterfly. 


Circumstances Surrounding the Description of M. millerorum 


The description of M. millerorum was published posthumously. Its 
description and associated type designations were complicated, there- 
fore, by the assembling of Clench’s unfinished manuscripts and the 
apparently related type series. This effort, made by Drs. Lee D. and 
Jacqueline Y. Miller was hampered in regard to millerorum by several 
circumstances. Firstly, Clench had not seen and had, therefore, not 
dissected the male paratype specimen from the American Museum of 
Natural History. Its existence had been called to his attention earlier 
by me. A description of the species had been prepared in manuscript 
by me, using the aforementioned specimen and a female in the British 
Museum (Natural History) collected in July at Bolanos, Jalisco State, 
Mexico. In 1979, I deferred description of the species to Clench, send- 
ing him information concerning both of the above specimens (the latter 
of which had also been listed by Shields, 1965, as M. spinetorum 
(Hewitson)). In turn, Clench forwarded for my examination the unique 
specimen from Otero County, New Mexico, noted by him (loc. cit., 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pp. 22-23) as collected “3 mi. N. Weed, ca. 6700 ft., 12 June 1977,” 
by H. K. and M. Clench. This specimen had the undersurface hindwing 
postbasal stripes characteristic of millerorum but was identified by 
Clench (loc. cit.) as an aberration of spinetorum. By the time of his 
death, Clench had not dissected the eventual holotype of millerorum, 
though I had returned my dissection of the Otero County specimen to 
him for comparison. 

When the Millers assembled the material for Clench (loc. cit.) none 
of the above material or information was located. Hence, when the 
original description was published, the Otero County specimen had 
still not been found, and although I possessed a drawing of its genitalia, 
I had not been able to compare it to either the eventual type or the 
Bolanos, Mexico, specimen. As a result there was no standard of com- 
parison to identify the Otero County specimen properly. Further, the 
male of millerorum remained undescribed; the treatment of millero- 
rum was ultimately limited to comments in the remaining unfinished 
manuscript of Clench, and information concerning millerorum cull- 
able from Shields (loc. cit.) was not integrated into the original descrip- 
tion. In subsequent years all of the above problems have been clarified. 


Clarifications Concerning M. millerorum, 
Its Biology and Occurrence 


Fig. 1 illustrates the features of the male and female genitalia of all 
known Arceuthobium (Loranthaceae)-feeding Mitoura taxa.' Shorter, 
hairlike spines on the cephalo-ventrad surface of the valvae (“‘bilobed 
configuration” sensu Johnson, 1976, 1978, and in press” are emphasized, 
while long hairlike spines not figured. The former, along with overall 
opaqueness in this valval region, characterize Arceuthobium-feeders 
contrasted to Cupressaceae-feeders (see Johnson, 1976, and in press) 
along with numerous other characters. Among Arceuthobium-feeders, 
the male genitalia of M. millerorum are distinctive as follows: (1) 
bilobed area markedly larger as contrasted to caudal length of valvae; 
(2) saccus widely parabolic; (3) cephalad cornutus at aedeagal terminus 
bifurcate; and (4) (not illustrated) caudal one-third of aedeagus dis- 
tinctly curved (60° angle) in known specimen. This latter trait occurs 
in no other known Mitoura though its generality in millerorum cannot 
be certain from the single known specimen. Differences in the female 
genital plate of Arceuthobium-feeders are most apparent in the nature 


1 The life histories of M. spinetorum and johnsoni (Skinner) have been well documented (see Shields, loc. cit.). That 
of millerorum is known only from the circumstantial data presented in this paper or inferred from the overall unity of 
morphological characters shared by taxa of Fig. 1 as compared to Mitoura taxa known to feed on Cupressaceae (see 
Johnson, 1976, 1978, and in press) [in press]. Revision of the Callophryina of the world with phylogenetic and biogeo- 
graphic analyses (Lepidoptera: Lycaenidae). Ms. (in part) as Ph.D. dissertation, Graduate Center, City University of 
New York (1981). 902 pp. 

*In press. Revision of the Callophryina of the world with phylogenetic and biogeographic analyses (Lepidoptera: 
Lycaenidae). Ms (in part) as Ph.D. dissertation, Graduate Center, City University of New York (1981). 902 pp. 


VOLUME 39, NUMBER 2 LDA! 





Fic. 1. Male and female genitalia of Arceuthobium-feeding Mitoura taxa—A, B, C: 
topotypical M. johnsoni male, ventral view of genitalia, caudal tip of aedeagus, and 
lateral view of valvae, respectively; D, E: topotypical M. johnsoni female, signum and 
genital plate, respectively. Using same display format—F, G, H (male); I, J (female): 
topotypical M. spinetorum; K, L, M (male): holotype M. estela; N, O (female): paratype 
M. estela; P, Q, R (male): paratype M. millerorum. Entry S (female): holotype M. 
millerorum; T, U (female): M. millerorum, Otero County, New Mexico. So as not to 
obscure other features, the male genitalia are drawn without the long, hairlike spines 
which protrude from the caudad portion of the valvae and ventrad surface of the uncus. 


of the lamellal lips caudad on the ductus bursae, the nature of the 
sclerotizations surrounding these lips, and the shape of the sculpturing 
caudo-ventrad on the ductus bursae. Readers familiar with genitalia 
of Cupressaceae-feeding Mitoura will note the overall differences ap- 
parent in these as compared to taxa in Fig. 1 (see Johnson, 1976, in 
press). Johnson (in press) attributes this hiatus to the relatively more 
primitive (plesiotypic) nature of the characters in genitalia of Ar- 
ceuthobium-feeders. Variation possible in the rather simplified struc- 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Undersurfaces of the wings of (A, left) M. millerorum, paratype male and 
(B, right) M. spinetorum (Fort Wingate, New Mexico, AMNH) chosen for its resem- 
blance to millerorum, having nearly all apparent wing characters except the postbasal 
stripes (see text for discussion). 


tural components of these is less than that in more highly specialized 
genitalic structures characteristic of Cupressaceae-feeders. As a result, 
relative significance of interspecific differences among taxa in Fig. 1 
must be viewed with reference to Mitoura taxa as a whole (particularly 
these seen as a primitive to derived hierarchy) rather than in terms of 
the few apparent differences within the Arceuthobium-feeders alone. 
With such a view, M. millerorum is clearly a sister species of spine- 
torum and not to be viewed as a subspecies. 

Only one of the illustrated species, M. estela (Clench), is not known 
to occur in the United States. In Fig. 1, item S illustrates the holotype 
of millerorum while T and U illustrate the Otero County, New Mexico, 
specimen. It is apparent from these comparisons (see figure explanation 
for details), along with the distinct wing characters mentioned by Clench 
and confirmed by my examination, that the latter represents millero- 
rum. The specimen has subsequently been located by Dr. John Rawlins 
at the Carnegie Museum of Natural History and recurated in that 
collection. Genitalic features of the Bolanos, Mexico, specimen are sim- 
ilar to both of the above mentioned specimens. 

Figure 2 illustrates the wing undersurfaces of the paratype male of 
millerorum (left, A; genitalia, Fig. 1 P-R) and a specimen of spine- 


VOLUME 39, NUMBER 2 123 


torum (right, B) selected for its overall resemblance to millerorum 
except for the postbasal stripes. The latter selection is relevant to 
Clench’s apparent reason for associating the Otero County specimen 
with spinetorum. Postbasal stripes in millerorum are usually more 
apparent than indicated by the photo and vary slightly from two dis- 
tinct slashes, generally in the same plane, to a single, long, straight 
stripe. The square shape of the distad extensions of the median line 
near the Thecla-spot also seems characteristic of millerorum as opposed 
to the more w-shaped distad extensions of the median line in spine- 
torum. Clench reasoned that the Otero County specimen was more 
like spinetorum in all characters except the postbasal stripes. Undoubt- 
edly this was because the limbal area of the hindwing undersurface on 
many spinetorum was like that on the Otero County specimen: bright- 
er and more flushed with gray-blue and with only small and obsoles- 
cent black spots along the submargin. The opposite tendency, however, 
also occurs as noted in Fig. 2. As a result it is important to summarize 
that the best overall superficial character for recognizing millerorum 
is its postbasal stripes. Comparatively, M. johnsoni is brown on the 
upper surface of the wings (not steel blue as on the remaining taxa in 
Fig. 1), while on estela the limbal area of the hindwing undersurface 
is accented by bright orange-red across the entire submargin (making 
it unmistakable). The association of M. dospassosi (Clench) with the 
above taxa (Clench, loc. cit.) was incorrect. It is clear from the mor- 
phology of dospassosi that it is most likely a Cupressaceae-feeder (see 
Johnson, in press). Clench’s assessment that the upper surface of dos- 
passosi appeared blue (Clench, loc. cit., p. 32: his apparent reason for 
the clustering) is questionable. Most specimens of dospassosi appear 
black or no more blue-tinted than specimens of M. sweadneri Cher- 
mock, a taxon with which dospassosi shares numerous genitalic char- 
acters and which feeds on Juniperus (Cupressaceae) (Johnson, 1978, 
in press). 

Clench (loc. cit.) mentioned the possibility that all Mexican speci- 
mens listed by Shields (loc. cit.) might represent millerorum. He was 
correct except, perhaps, for the specimen from Baja California Norte. 
Except for the latter, all of the above specimens have been seen by me 
and represent millerorum. The specimen from Baja California (for 
which Patterson & Powell, 1959, mention no unusual features) is well 
within the zoogeographical region characterized by the distribution of 
spinetorum (and the tendency of numerous populations of butterflies 
in southern California to have associations southward into northern 
Baja). A larval specimen of Mitoura, perhaps referable to millerorum 
and located in the Peabody Museum (Yale University), is of great 
interest. It is from “6 mi. E. of the Mexico-Michoacan boundary on 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hi. 45, leg. Hawksworth and Wiens” (Shields, loc. cit., and Peabody 
Museum). No adult was reared from this larva and, therefore, its spe- 
cific identity as either of the Mexican taxa millerorum or estela cannot 
be ascertained with certainty. However, it was collected on Arceutho- 
bium globosum Hawksworth and Wiens growing on Pinus michoa- 
cana Martinez (Pinaceae) and is identifiable as a Mitoura. If the spec- 
imen represents either of the taxa millerorum or estela, this apparent 
unique monophagy would be a character equivalent to that which, 
among others, distinguishes the specificity of M. johnsoni (considered 
monophagous on A. douglassi Engelm., Shields, loc. cit.; Johnson, 1976, 
and in press). 
Conclusions 


Four specimens are presently known of M. millerorum, three from 
Mexico and one from the United States. A larval specimen may rep- 
resent the species and provides possible clues concerning its biology. 
The species distribution, relative to congeners and other related Eu- 
maeini lycaenids, is characteristic of segregations occurring from mon- 
tane central Mexico north to the isolated mountain ranges of southern 
Arizona and New Mexico (Johnson, in press; Rosen, 1975). This, along 
with its relative ease of identification, makes it a taxon which should 
be pursued by North American lepidopterists, particularly in regard 
to elucidation of its biology. As is well known, all of the hitherto men- 
tioned species of Mitoura occur in highly local populations of low 
density (Brown, Eff & Rotger, 1957; Ferris & Brown, 1981). In the 
case of millerorum, however, its occurrence in the southwestern United 
States demonstrates that one should not assume that spinetorum will 
be the only species collectable by concerted fieldwork. 


LITERATURE CITED 


BROWN, F. M., D. EFF & B. ROTGER. 1957. Colorado butterflies. Denver Museum Nat. 
Hist., Denver. vii + 368 pp., 314 figs. 

CLENCH, H. K. 1981. New Callophrys (Lycaenidae) from North and Middle America. 
Bull. Alynn Mus. 64:1-31. 

FERRIS, C. D. & F. M. BRown. 1981. Butterflies of the Rocky Mountain states. Uni- 
versity of Oklahoma Press, Norman. xviii + 442 pp., 26 figs., 314 maps. 

JOHNSON, K. 1976. Three new Nearctic species of Callophrys (Mitoura), with a diag- 
nostis [sic] of all Nearctic consubgeners (Lepidoptera, Lycaenidae). Bull. Allyn Mus. 
38:1-30. 

1978. Specificity, geographic distribution, and foodplant diversity in four Cal- 
lophrys (Mitoura) (Lycaenidae). J. Lepid. Soc. 32:3-19. 

PATTERSON, D. & J. A. POWELL. 1959. Lepidoptera collecting in the Sierra San Pedro 
Martir, Baja California. J. Lepid. Soc. 13:229-235. 

ROSEN, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24:431- 
464. 

SHIELDS, O. 1965. Callophrys (Mitoura) spinetorum and C. (M.) johnsoni: Their 
known range, habits, variation and history. J. Res. Lepid. 4:233-250. 





Journal of the Lepidopterists’ Society 
39(2), 1985, 125-133 


TECHNIQUES FOR MAINTAINING A CULTURE OF THE 
BLACK SWALLOWTAIL BUTTERFLY, 
PAPILIO POLYXENES ASTERIUS STOLL (PAPILIONIDAE) 


MAUREEN CARTER AND PAUL FEENY 


Section of Ecology and Systematics, Cornell University, 
Ithaca, New York 14853 


ABSTRACT. A culture of the black swallowtail butterfly, Papilio polyxenes asterius 
Stoll (Papilionidae) is initiated from field-collected females. One hundred to two hundred 
larvae are reared on potted plants in a greenhouse. Adults are housed in an environmental 
growth chamber with a 16 L/8 D photoperiod, day and night temperatures of 27°C and 
15.5°C, respectively, and with a relative humidity at 70 + 15%. Adults are hand-fed and 
hand-paired. 


Successful rearing techniques are often a prerequisite for experi- 
mental success, yet they are seldom discussed in scientific articles. Our 
study of the behavior of the black swallowtail butterfly (Papilio po- 
lyxenes asterius Stoll) has required the maintenance of a year-round 
culture of this butterfly. Here we describe techniques that may be 
useful to other researchers who would like to rear this or related but- 
terfly species. We do not claim originality for many of these tech- 
niques; some are scattered in the literature, while others have been 
developed by staff and graduate students in our research group or 
suggested to us by lepidopterists elsewhere. 


Initiating a Culture 


Collecting. A stock of wild female butterflies is collected locally. In 
central New York, peak collecting is generally during the second an- 
nual brood, from early July to late August (Lederhouse, 1978). Led- 
erhouse (1981) found only 2.3% of field-caught females to be virgins, 
and we assume that our wild-collected females have already been fer- 
tilized. Captured butterflies are placed in 3%” square, glazed paper 
envelopes! and transported back to the laboratory in a cool, shady place 
in the vehicle. In the laboratory, they are housed in a walk-in environ- 
mental growth chamber? under a 16 L/8 D photoperiod, day and night 
temperatures of 27°C and 15.5°C, respectively, and at a relative hu- 
midity of 70 + 15%. 

Feeding. Butterflies are immediately fed a 10% solution of honey in 
water, poured into the inverted top of a small petri dish, the inverted 
bottom of which floats on the solution (Fig. 1). Each butterfly is held 
close to the dish and its proboscis is unrolled with an insect pin; the 


1 Ward's Natural Science Establishment Inc., P.O. Box 92912, 5100 West Henrietta Road, Rochester, NY 14692-9012. 
2 Environmental Growth Chambers, P.O. Box 407, Chagrin Falls, OH 44022. 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1 & 2. 1, butterflies at honey-water feeding station; 2, undersides of butterfly 
wings illustrating markings for number 68. 


tip of the proboscis is placed into the honey-water. If feeding occurs, 
it will continue for 1-5 minutes. This technique allows butterflies to 
feed with minimal contamination of the legs, abdomens and wings. 
We have been unable to persuade P. polyxenes adults to initiate feed- 
ing in the laboratory without assistance. 

Numbering. We use a 1-2-4-7 marking system (Ehrlich & Davidson, 
1960; Brussard, 1971; Southwood, 1978) as illustrated in Fig. 2. This 
technique was first used in our black swallowtail culture by Lederhouse 
(1978). Butterflies can be numbered from | to 99; additional numbers 
are available by changing marker color. | 

Oviposition. After numbering and feeding, each female butterfly is 
caged with a potted host plant. Cages are wood-framed, 34 cm X 46 
cm, and covered on five sides with organdy mesh or “no-see-um”’ 
netting. We do not use wire mesh, because the resistance produced by 
gripping butterfly tarsi can cause them to break off in handling. Old 
or worn specimens are placed in plastic shoe boxes? lined with paper 
towels and containing sprigs of host plant fitted with water-filled 
“Aquapics’’®,* as shown in Fig. 8. Lids are left ajar or the center is cut 
out and replaced with wire mesh for ventilation. Because host plant 
sprigs will desiccate before eggs hatch, the eggs are removed by light 
nudging with a fingernail and are placed in a plastic petri dish,® 15 cm 
in diameter, lined with moist filter paper. 


5 Tri-State Molded Plastics, Inc., P.O. Box 6, Dixon, KY 42409. 
“Cleveland Plant and Flower Co., Wholesale Florists, 262-272 Clinton Street, Binghamton, NY 13905. 
5 VWR Scientific Inc., P.O. Box 1050, Rochester, NY 14603. 


VOLUME 39, NUMBER 2 |g 





Fics. 3-7. 3, plastic shoe box with host plant sprigs in “Aquapics’® for oviposition 
or larval feeding; 4, individual larval rearing cage; 5, butterfly emergence cage; 6, hand- 
pairing of butterflies; 7, butterflies in copula. 


Rearing Immature Stages 


We have relied on two methods of rearing larvae: (1) on potted 
plants, and (2) on excised leaves in closed containers. 

Rearing on potted plants. Egg-laden plants are kept on a cart in the 
greenhouse, under a 16-hour light cycle provided by 400 and 1000 
watt metal halide lamps.° Egg-laden plants are watered daily. De- 
pending on temperature, eggs hatch in 3-5 days. In the greenhouse, 
when larvae are present, temperatures are maintained between 24 and 
27°C. Automatic misting systems should not be used for watering plants 
containing larvae younger than the fourth instar, due to high mortality 
from drowning. 

As larvae grow, fresh pots of food plant are placed around them. 
The larvae will remain on the plants if an ample supply of food is 
available. Frass is swept out daily. Because larval feeding is minimal 


6 General Electric Company, Hendersonville, NC. 


128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


during the first three instars, an abundance of larvae can be main- 
tained. At the fourth instar, the number of larvae is decreased to the 
number of adults needed, plus 15% to allow for accidental deaths and 
abnormal specimens. Culture larvae are transferred directly to potted 
food plants on a greenhouse bench, where they range freely until the 
end of the fifth instar. If unconfined, they will wander extensively to 
find a pupation site. To prevent such larval wandering, we confine the 
late fifth instar larvae with their potted food plants under metal-framed 
wire cages, 71 cm x 71 cm, or under cylinders of wire with organdy 
tops that fit over a single pot (Fig. 4). After completing their feeding 
and voiding their guts, larvae pupate on the roof or walls of the cage 
and can easily be collected for transfer to emergence cages. 

When fresh food plants are available in the field, larvae are reared 
on potted plants in the greenhouse until the fourth instar and then on 
excised stems placed in water-filled jars or Erlenmeyer flasks. Food 
plant is changed every second day, but water is changed daily. Wire 
cages cover these containers. 

Rearing in closed containers. Larvae can be reared in different 
types of closed containers. Eggs are removed from plants and trans- 
ferred to petri dishes kept in our growth chamber. With a fine camel- 
hair brush or a broken boiling stick, newly-hatched first-instar larvae 
are moved to excised sprigs of food plant fitted with “Aquapics’’® 
(Scriber, 1977). The food plant is placed in a plastic shoe box lined 
with paper towels. Because of heavy condensation, paper towels must 
be changed daily and boxes wiped out. Individual larvae can also be 
reared in small glass petri dishes or plastic containers with tight-fitting 
lids and moist filter-paper bottoms. Cut leaves are placed in these 
without any reservoir. The idea is to keep the leaves from drying out 
without drowning the larvae. Small rearing dishes must be wiped out 
every 1-2 days. This method is used frequently by researchers feeding 
weighed or treated leaf material. Food plant in closed containers can 
be consumed for two days, but water reservoirs must be filled and 
liners misted daily. A small twig added to the container is a preferred 
pupation site. Containers are sterilized routinely in 5% sodium hypo- 
chlorite solution. Rearing a large number of larvae in closed containers 
is extremely time-consuming. 

Pupae. Larvae pupate on plant stems and pots or the frame or wire 
of cages. Prepupae shed their ultimate larval skin in 24-48 hours. 
Prepupae and new pupae are easily damaged. They are removed from 
pupation sites by misting the silk pad with water and pulling gently 
at the silk. Pupae are moved to emergence cages in the growth cham- 
ber, 5-6 per cage. Each emergence cage is a plastic petri dish, with 


VOLUME 89, NUMBER 2 129 





Fic. 8. Abdominal apices of female and male pupae (GO, genital opening; S8, seg- 
ment 8; S9-10, segments 9 and 10; AO, anal opening). 


a “Kimwipe’® liner on the bottom and a 6” cylinder of wire mesh 
(Fig. 5). 

During June a group of larvae is reared under a short day photo- 
period (8 L/16 D) to obtain diapausing pupae. They are stored in a 
cold environment (0°C) until needed to initiate a new culture, if nec- 
essary, during the winter. 

Sex of pupae can be accurately determined by close examination of 
the sutures on the genital plates (Jackson, 1890; Poulton, 1890; Mosher, 
1916). The male genital opening is on the mid-ventral surface of seg- 
ments 9+10 (Fig. 8, GO); while in the female, two genital openings 
are confluent forming a single slit across the boundaries of segments 8 
and 9+ 10. The genital opening is surrounded by a raised area in both 
sexes (but more prominently in the female). 


Adults 


In the laboratory, the adult emergence pattern is protandrous, with 
eclosion following pupation by 138-20 days (Lederhouse et al., 1982). 
Eclosing butterflies dry for 24 hours. One day after emergence, each 
female is marked, fed and bred by the hand-pairing technique devel- 
oped by Clarke and Sheppard (1956) as illustrated in Fig. 6. Males are 
fed but not marked. Because Sims (1979) found spermatozoa counts to 
be low in young males of a closely related species, Papilio zelicaon 
Lucas, males are not mated for 48 hours after eclosing. Each mated 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pair of butterflies (Fig. 7) is placed on the inside of a tilted cage. They 
crawl to the top and hang for 51.3 + 8.6 minutes (Lederhouse, 1981). 
Male butterflies can be discarded after the initial mating. If not, they 
can be remated if rested 48 hours, but spermatophore size will probably 
be decreased (see Sims, 1979) and copulation time will be longer (Led- 
erhouse, 1981). When the majority of eclosing females have mated, 
new pots of host plants are introduced to collect 300-400 eggs for the 
next generation of the culture. Additional heat has been provided to 
increase oviposition activity. Sixty-watt incandescent or infrared re- 
flector lamps are suspended a minimum of 46 cm above the cages for 
a period of 3-4 hours. Because egg fertility decreases with time from 
initial mating (Lederhouse, 1981), we discard mated females after 7- 
10 days. 

During periods of absence, pupae and adults can be stored. Pupae 
are packed in moist paper towels (see Stone & Midwinter, 1975) and 
placed in a small insulated cooler. The cooler is stored in a cold envi- 
ronment (0°C) which halts adult emergence. They can stay there for 
up to two weeks. Adults can be safely refrigerated for 2-3 days. They 
are put into glazed paper envelopes and placed upright in a cooler 
after feeding. Butterflies emerging that day are allowed to dry until 
late afternoon before storage. After removal from the cold environ- 
ment, they need 2-3 hours to warm up before feeding, the first hour 
in the envelopes. 

Unfortunately, larvae cannot go unattended, even for 24 hours. Pot- 
ted plants with larvae feeding on them must be watered daily. 


Providing Food Plant 


Greenhouse plants. Prerequisites for maintaining a culture of this 
butterfly are a large amount of greenhouse space, readily available 
food plant in the field, or some combination of the two. We need 325 
square feet of -greenhouse bench space, supplemented by field-collect- 
ed plants from June through August, to produce two broods of adults 
(100-200 individuals each) emerging every six-week period. 

Our supply of greenhouse plants for each coming year is started 
from seed in the spring. Seeds are sown 10-20 per 6” pot, in sterilized 
soil or artificial mix. We routinely plant seeds of carrot (Daucus carota 
L.) and parsley (Petroselinum crispum (Mill.) Mansfeld). Other food 
plant species can be planted from commercial or field-collected seeds 
(Tietz, 1972; Rehr, 1973; Tyler, 1975; Berenbaum, 1978; Scriber & 
Finke, 1978). Seedlings emerge in two weeks and are thinned before 
plants are ready for use two months later. Because these plant species 
are cold weather crops, temperatures in the greenhouse (when larvae 


VOLUME 39, NUMBER 2 131 


are not present) are kept at 21°C. Evaporative cooling units keep green- 
house temperatures down in the summer. 

Plants are routinely watered and fertilized, depending on soil type 
and climatic conditions. Fertilizing is especially important to promote 
new growth after the stress of larval feeding. The metal halide lamps, 
providing a 16-hour photoperiod for larvae, also enhance plant growth 
in winter. All plants are treated to combat the usual greenhouse pests. 
We avoid broad-spectrum insecticides and use only those specific for 
target pest organisms. 

After larval feeding, plant stems are cut short to facilitate new growth 
and provide optimal coverage for pesticide applications. At any one 
time, % of our plants are usable for feeding, 4% are waiting out twice 
the residual time of the last pesticide application and % are just starting 
to put out new growth. Destruction of some plants by larvae, pruning 
and pest attacks necessitate having more than one plant per pot. New 
plants are potted each year, because plants carried over from one year 
to the next send out flowering stalks and regenerate little new leaf 
material. 

Field-collected leaves. To support larvae in the summer when new 
greenhouse plants are still maturing, cuttings are collected from wild 
plants in the field or from plants cultivated in a garden plot. Cuttings 
are taken in early morning. Cut stems are placed in a bucket of water 
that is surrounded by ice in a cooler. By taking these precautions, 
cuttings need only be changed every other day. Cuttings taken during 
the heat of the day and not adequately protected from dehydration do 
not remain turgid. New leaves, flower and seed heads are collected. 
New growth of biennial species can be found in spring and again in 
late summer. 


Morbidity and Mortality 


Disease suppression in insects is dependent upon the maintenance 
of optimal living and rearing conditions (Steinhaus, 1963). These op- 
timal conditions include freedom from stress such as that induced by 
crowding, toxic chemicals, adverse conditions of light and radiation, 
inadequate nutrition or lack of oxygen (Steinhaus, 1958, 1963; Burges, 
1973). Our rearing method, using potted plants in the greenhouse, 
provides maximum radiation, air circulation, humidity and space, and 
optimal food plant quality. Although growth chambers allow more 
precise control of temperature, humidity and light cycle fluctuations, 
the incidence of disease in such environments is higher. 

Our culture techniques have eliminated epizootic infections to im- 
mature stages. However, some enzootic diseases are encountered, and 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


as Burges (1973) points out most species of leaf-feeding Lepidoptera 
probably possess a nuclear polyhedrosis virus, a cytoplasmic polyhe- 
drosis virus, a granulosis virus and microsporidians. The small per- 
centage of mortality seen every culture cycle is symptomatic of a fatal 
bacterial septicemia (Bucher, 1960). Affected larvae discontinue to feed 
or grow and eventually die; dead larvae hang limp and flaccid, a 
certain sign of extensive tissue destruction and putrefaction of body 
contents. The invading bacteria are probably in a class defined by 
Bucher (1960) as “‘potential pathogens.’ These pathogens invade and 
multiply in the susceptible hemocoele after a variety of stress factors 
has made the gut more permeable. 

No epizootic infections have occurred in the imagos of our culture, 
although we do see some malformations such as missing or shortened 
appendages, deformed tarsi, deformed claspers of males, and very small 
or oversized individuals. One environmental aberration that was once 
a persistent problem is the condition of a split proboscis. Providing 
high humidity in the local environment of pupae, either by resting 
them on a damp substrate (see Stone & Midwinter, 1975) or misting 
daily (Lederhouse, 1978), can minimize its occurrence. All malformed 
pupae and adults are eliminated routinely from the culture. 

From June to October, prepupae and pupae in the greenhouse must 
be protected from attack by the hymenopterous parasitoid Pteromalus 
puparum (L.) (Pteromalidae). Cages can be covered with “‘no-see-um”’ 
netting to keep these parasites out. 


Maintaining Genetic Variability 


Heterozygosity in a domesticated laboratory culture can be lost 
quickly, especially if the culture is initiated from a few individuals 
(Benz, 1963). An increase in homozygosity is believed to contribute to 
a decline in fitness. To maintain genetic variability, we collect many 
female butterflies from the local wild population to initiate a new 
culture or add genetic material to an existing one. To avoid inbreeding 
depression (Mayr, 1970), during the winter months we rear two groups 
of larvae every six weeks, each with 100-200 individuals. This gives 
us 50-100 mated females to generate the next cycle. 


ACKNOWLEDGMENTS 


We thank E. Richard Hoebeke, J. Mark Scriber, May Berenbaum, Mark Evans and 
Lorraine Rosenberry for their suggestions for improving this paper. Photographs 3, 4, 5, 
6 and 7 were taken by E. Richard Hoebeke. Jim Miller provided Fig. 8. We are grateful 
to Lorraine Rosenberry for her valuable input in the development of many of these 
techniques. The National Science Foundation has provided continuing support for this 
work. Mention of a trademark or a proprietary product does not constitute a guarantee 
or a warranty of the product by the National Science Foundation or Cornell University, 
nor imply its approval to the exclusion of other products that also may be suitable. 


VOLUME 39, NUMBER 2 133 


LITERATURE CITED 


BENZ, G. 1968. Genetic diseases and aberrations. In Insect pathology (E. A. Steinhaus, 
ed.), pp. 161-189. Academic Press, New York. 

BERENBAUM, M. 1978. Taenidia integerrima, a new foodplant record for Papilio po- 
lyxenes (Papilionidae). J. Lepid. Soc. 32(4):303-304. 

BLAU, W. S. 1981. Life history variation in the black swallowtail butterfly. Oecologia 
48:116-122. 

BRUSSARD, P. F. 1971. Field techniques for investigations of population structure in a 
“ubiquitous” butterfly. J. Lepid. Soc. 25(1):22-29. 

BUCHER, G. E. 1960. Potential bacterial pathogens of insects and their characteristics. 
J. Insect Pathol. 2:172-195. 

BurGEs, H. D. 1973. Enzootic diseases of insects. Ann. N.Y. Acad. Sci. 217:31-49. 

CLARKE, C. A. & P. M. SHEPPARD. 1956. Hand-pairing of butterflies. Lepid. News 
10(1-2):47-53. 

EHRLICH, P. R. & S. E. DAVIDSON. 1960. Techniques for capture-recapture studies of 
Lepidoptera populations. J. Lepid. Soc. 14(4):227-229. 

JACKSON, W. H. 1890. Studies in the morphology of the Lepidoptera. Part I. Linn. Soc. 
London, Trans., Ser. 2, Zool. 5:143-186. 

LEDERHOUSE, R. C. 1978. Territorial behavior and reproductive ecology of the black 
swallowtail butterfly, Papilio polyxenes asterius Stoll. Unpublished Ph.D. thesis, Cor- 
nell University, Ithaca, New York. 155 pp. 

1981. The effect of female mating frequency on egg fertility in the black 
swallowtail, Papilio polyxenes asterius (Papilionidae). J. Lepid. Soc. 35(4):266-277. 

LEDERHOUSE, R. C., M. D. FINKE & J. M. SCRIBER. 1982. The contributions of larval 
growth and pupal duration to protandry in the black swallowtail butterfly, Papilio 
polyxenes. Oecologia 53:296-300. 

Mayr, E. 1970. Populations, species, and evolution. Belknap Press, Cambridge. 453 pp. 

MOosHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa. 
Bull. Illinois State Lab. Nat. Hist. 12(2):14-159. 

POULTON, E. B. 1890. The external morphology of the lepidopterous pupa: Its relation 
to that of the other stages and to the origin and history of metamorphosis. Parts I- 
III. Linn. Soc. London, Trans., Ser. 2, Zool. 5:187-212. 

REHR, S. S. 1973. New foodplant records for Papilio polyxenes F. (Papilionidae). J. 
Lepid. Soc. 27(3):237-238. 

SCRIBER, J. M. 1977. Limiting effects of low leaf-water content on the nitrogen utili- 
zation, energy budget, and larval growth of Hylaophora cecropia (Lepidoptera: 
Saturniidae). Oecologia 28:269-287. 

SCRIBER, J. M. & M. FINKE. 1978. New foodplant and oviposition records for the eastern 
black swallowtail, Papilio polyxenes on an introduced and a native umbellifer. J. 
Lepid. Soc. 32(3):236-238. 

Sims, S. R. 1979. Aspects of mating frequency and reproductive maturity in Papilio 
zelicaon. Amer. Midl. Natur. 102(1):36-50. 

SOUTHWOOD, T. R. E. 1978. Ecological methods. Chapman and Hall, London. 524 pp. 

STEINHAUS, E. A. 1958. Stress as a factor in insect disease. Proc. 10th Int. Congr. 
Entomol. 4:725-730. 

1963. Introduction. In Insect pathology (E. A. Steinhaus, ed.), pp. 1-27. Aca- 
demic Press, New York. 

STONE, J. L. S. & H. J. MIDWINTER. 1975. Butterfly culture. Blandford Press Ltd., 
Dorset. 104 pp. 

TiETZ, H. M. 1972. An index to the described life histories, early stages and hosts of 
the Macrolepidoptera of the continental United States and Canada, Vol. 1. A. C. 
Allyn, Sarasota. 536 pp. 

TyLerR, H. A. 1975. The swallowtail butterflies of North America. Naturegraph, Healds- 
burg. 192 pp. 








Journal of the Lepidopterists’ Society 
89(2), 1985, 134-138 


THE BUTTERFLIES OF MISSISSIPPI— 
SUPPLEMENT NO. 3? 


BRYANT MATHER? AND KATHARINE MATHER 
213 Mt. Salus Road, Clinton, Mississippi 39056 


ABSTRACT. An annotated list of Mississippi butterflies is presented. This updated 
version is the fifth such list published. Six names additional to the previous lists have 
been included. 


Of the five published lists of Mississippi butterflies, this is the first to 
use the names and arrangement of Miller and Brown (1981) as amend- 
ed by them in Hodges (Editor) (1983). It includes six names not in- 
cluded in the fourth list (Mather & Mather, 1976). The growth rate 
has dropped to fewer than one per year as indicated in Table 1 below: 


TABLE 1. Published lists of Mississippi butterflies, showing rate of increase in the 
addition of names previously unrecorded from the state. 


Time Names added 
List Reference Names interval Names added per year 
1 Weed (1894) 53 — —_ — 
2 Hutchins (1933) 78 39 20 0.5 
3 M. & M. (1958) 122 25 49 1:9 
4 M. & M. (1976) 144 18 22 1.2 
5 M. & M. (1984) 150 8 6 0.75 


In 1958 we expressed the opinion that the list would grow to include 
about 160 names. We also said, “‘there may be cases in which the 
Mississippi representatives of a given species represent more than one 
population; if so, we do not believe that we as yet have adequate data 
to support such a conclusion.” Now we do. Our reasons for adding the 
six names are summarized below. 

1. Papilio glaucus australis Maynard. In 1958, we noted that Pro- 
fessor R. L. Chermock had told us that the south Alabama glaucus 
population was referable to australis. We now have sufficient material 
from south Mississippi to come to the same conclusion about the pop- 
ulation there. 

2. Papilio troilus ilioneus J. E. Smith. In 1958, on the advice of Mr. 
Cyril F. dos Passos, we referred the Mississippi population to the sub- 
species. Before 1976, Mr. Harry K. Clench examined a long series and 
concluded that the correct referral was to the nominate subspecies. We 


‘Contribution No. 589, Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and 
Consumer Services, Gainesville, Florida 32602. 
* Research Associate, Florida State Collection of Arthropods. 


VOLUME 39, NUMBER 2 135 


therefore used that name in the 1976 list. We are now of the opinion 
that both subspecies are present. This view tends to be supported by 
the treatment of texanus Ehrmann, type locality Houston, Texas as a 
synonym of ilioneus J. E. Smith, type locality Georgia. 

3. Basilarchia archippus floridensis (Strecker). In 1958 we assigned 
the Mississippi population to B. a. watsoni (dos Passos) on the advice 
of Mr. dos Passos. We also noted that a specimen from Harrison Co. 
in the collection at the University of Missouri at Columbia had been 
determined as B. a. floridensis by Harold I. O’Byrne, while another, 
from Tishomingo Co., in the Los Angeles County Museum had been 
determined as B. a. archippus by J. A. Comstock. We now believe that 
both B. a. watsoni and B. a. floridensis populations are present in 
Mississippi. | 

4. Asterocampa alicia (W. H. Edwards). In 1958 we assigned the 
Mississippi population to A. celtis alicia, type locality New Orleans, 
La., on the advice of Mr. dos Passos, in spite of the fact it was recog- 
nized by him that the majority of the specimens in the sample he had 
seen resembled A. c. celtis, type locality Georgia. In 1975, Howe ele- 
vated alicia to species status based on unpublished work by W. J. 
Reinthal. If celtis and alicia are not conspecific, then the population 
in Mississippi nearest to New Orleans must be assigned to alicia and 
the remainder (superficially indistinguishable from celtis) assigned to it. 

5. Asterocampa flora (W. H. Edwards). Although in 1958 our Mis- 
sissippi sample included material resembling flora, type locality Pa- 
latka, Florida, we called the population A. c. clyton, type locality Geor- 
gia, on the advice of Mr. dos Passos, who expressed the view that the 
name flora should be restricted to the Florida population. Howe (1975) 
also elevated flora to species status, based on unpublished work by 
Reinthal and gave its distribution as “Southern Georgia, Florida, Gulf 
States, and Texas.” We thus assign the material that resembles flora 
(coming from the southern third of Mississippi) to flora and that re- 
sembling clyton (coming from north of there) to clyton. 

6. Cercyonis pegala alope (Fabricius). In 1958, only a few records 
were known, primarily from the southern part of the state. In 1976, it 
was clear that this material belonged to the subspecies named abbotti 
by F. M. Brown in 1969. In 1979, David Hess examined all available 
Mississippi material and determined a series from northern Mississippi 
(Tippah Co.) as C. p. alope, while those from the southern part of the 
state remained C. p. abbotti. 

The revised check list, including the six additional names, using the 
names and sequence as given by Miller and Brown (1981) as amended 
by them in Hodges et al. (1983) follows: 


136 


O OID TWP be 


. 3870. 


3886. 
3889. 
3902. 
3904. 
3909. 


. 3910. 
. 3913. 
. 3932. 
. 3946. 
. 3947. 
. 3952. 
, 3954. 
. 3956. 
. 3957. 
. 3959. 
. 3966. 
. 3968. 
» Beil Te 
. 3993. 
. 3995. 
. 3998. 
. 4004. 
. 4010. 
. 4013. 
. 4027a. 
. 4029. 
. 4041. 
. 4042. 
. 4045. 
. 4046. 
. 4047. 
. 4048. 
. 4049a. 
. 4050. 
. 4051. 
. 4052. 
. 4059. 
. 4060. 
. 4063. 
. 4064a. 
. 4070. 
. 4071. 
. 4078. 
. 4074. 
. 4078a 
. 4080. 
. 4084. 
. 4096. 
. 4099. 
. 4100. 
. 4101. 
. 4105. 
. 4107. 
. 4108. 
. 4111. 
. 4114. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Epargyreus clarus clarus (Cramer) 

Urbanus proteus (Linnaeus) 

Urbanus dorantes dorantes (Stoll) 
Autochton cellus (Boisduval and Leconte) 
Achalarus lyciades (Geyer) 

Thorybes bathyllus (J. E. Smith) 

Thorybes pylades (Scudder) 

Thorybes confusis Bell 

Staphylus hayhurstii (W. H. Edwards) 
Erynnis brizo brizo (Boisduval and Leconte) 
Erynnis juvenalis juvenalis (Fabricius) 
Erynnis horatius (Scudder and Burgess) 
Erynnis martialis (Scudder) 

Erynnis zarucco (Lucas) 

Erynnis funeralis (Scudder and Burgess) 
Erynnis baptisiae (Forbes) 

Pyrgus communis (Grote) 

Pyrgus oileus (Linnaeus) 

Pholisora catullus (Fabricius) 

Nastra lherminier (Latreille) 

Nastra neamathla (Skinner and R. C. Williams) 
Lerema accius (J. E. Smith) 

Ancyloxypha numitor (Fabricius) 
Copaeodes minimus (W. H. Edwards) 
Hylephila phyleus (Drury) 

Hesperia metea licinus (W. H. Edwards) 
Hesperia attalus attalus (W. H. Edwards) 
Polites themistocles (Latreille) 

Polites origenes origenes (Fabricius) 

Polites vibex vibex (Geyer) 

Wallengrenia otho (J. E. Smith) 
Wallengrenia egeremet (Scudder) 
Pompeius verna (W. H. Edwards) 
Atalopedes campestris huron (W. H. Edwards) 
Atrytone arogos arogos (Boisduval and Leconte) 
Atrytone delaware delaware (W. H. Edwards) 
Problema byssus byssus (W. H. Edwards) 
Poanes hobomok (Harris) 

Poanes zabulon (Boisduval and Leconte) 
Poanes yehl (Skinner) 

Poanes viator zizaniae Shapiro 

Euphyes arpa (Boisduval and Leconte) 
Euphyes pilatka (W. H. Edwards) 

Euphyes alabamae (Lindsey) 

Euphyes dukesi (Lindsey) 

Euphyes ruricola metacomet (Harris) 
Atrytonopsis hianna hianna (Scudder) 
Atrytonopsis loammi (Whitney) 
Amblyscirtes hegon (Scudder) 

Amblyscirtes aesculapius (Fabricius) 
Amblyscirtes carolina (Skinner) 
Amblyscirtes reversa (F. M. Jones) 
Amblyscirtes vialis (W. H. Edwards) 
Amblyscirtes belli H. A. Freeman 
Amblyscirtes alternata (Grote and Robinson) 
Lerodea eufala (W. H. Edwards) 

Oligoria maculata (W. H. Edwards) 


VOLUME 39, NUMBER 2 


AS: 
AL 16. 
. 4119. 
5 Gta SY 
ALT. 
. 4159a. 
4170. 
44/6. 
. 4176b. 
. 4181. 
. 418 1a. 
. 4182. 
. 4184. 
. 4198. 
. 4197. 
. 4198a. 
. 4207. 
. 4209. 
. 4210. 
. 4224. 
S422 7a. 
. 4228a. 
. 4229. 
) 4D8Ye 
aoa 
» 4243: 
. 4246. 
. 4248. 
. 4249. 
. 4256. 
. 4268a 
nA 0: 
_ 4275a. 
- 4282a. 
. 4284. 
. 4285a. 
. 4299. 
. 4300. 
. 4318. 
. 4822b. 
. 43826c. 
. 4328. 
m4 3o2: 
. 4335. 
. 4336. 
. 4354a. 
RrASbi. 
. 4359a. 
. 43860a. 
. 4361. 
. 4868. 
. 4386. 
. 4410. 
. 4418a. 
. 4418a. 
. 4420. 
» GA Olle 


Calpodes ethlius (Stoll) 

Panoquina panoquin (Scudder) 

Panoquina ocola (W. H. Edwards) 
Megathymus yuccae yuccae (Boisduval and Leconte) 
Battus philenor philenor (Linnaeus) 

Papilio polyxenes asterius Stoll 

Papilio cresphontes cresphontes Cramer 
Papilio glaucus glaucus Linnaeus 

Papilio glaucus australis Maynard 

Papilio troilus troilus Linnaeus 

Papilio troilus ilioneus J. E. Smith 

Papilio palamedes Drury 

Eurytides marcellus Cramer 

Pontia protodice (Boisduval and Leconte) 
Artogeia rapae (Linnaeus) 

Ascia monuste phileta (Fabricius) 
Falcapica midea midea (Hiibner) 

Colias philodice philodice Godart 

Colias eurytheme Boisduval 

Zerene cesonia (Stoll) 

Anteos maerula lacordairei (Boisduval) 
Phoebis sennae eubule (Linnaeus) 

Phoebis philea (Johansson) 

Eurema lisa Boisduval and Leconte 
Eurema nicippe (Cramer) 

Eurema daira (Godart) 

Eurema mexicanum (Boisduval) 

Nathalis iole Boisduval 

Feniseca tarquinius tarquinius (Fabricius) 
Hyllolycaena hyllus (Cramer) 

Eumaeus atala florida Réber 

Atlides halesus halesus (Cramer) 
Harkenclenus titus mopsus (Hiibner) 
Satyrium calanus falacer (Godart) 
Satyrium kingi (Klots and Clench) 
Satyrium liparops strigosum (Harris) 
Calycopis cecrops (Fabricius) 

Calycopis isobeon (Butler and Druce) 
Mitoura grynea grynea (Hiibner) 

Incisalia augustus croesioides (Scudder) 
Incisalia henrici turneri Clench 

Incisalia niphon niphon (Hubner) 
Eurystrymon ontario ontario (W. H. Edwards) 
Parrhasius m-album (Boisduval and Leconte) 
Strymon melinus melinus Hiibner 
Brephidium isophthalma pseudofea (Morrison) 
Leptotes marina (Reakirt) 

Hemiargus ceraunus antibubastus Hiibner 
Hemiargus isola alce (W. H. Edwards) 
Everes comyntas comyntas (Godart) 
Celastrina ladon ladon (Cramer) 

Calephelis virginiensis (Guérin-Méneville) 
Libytheana bachmanii bachmanii (Kirtland) 
Agraulis vanillae nigrior Michener 
Heliconius charitonius tuckeri W. P. Comstock and F. M. Brown 
Polygonia interrogationis (Fabricius) 
Polygonia comma (Harris) 


137 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


115. 4482. Nymphalis antiopa antiopa (Linnaeus) 
116. 4434. Vanessa virginiensis (Drury) 

117. 4435. Vanessa cardui (Linnaeus) 

118. 4437a. Vanessa atalanta rubria (Fruhstorfer) 

119. 4439. Hypolimnas misippus (Linnaeus) 

120. 4440. Junonia coenia (Hiibner) 

121. 4443a. Anartia jatrophae guantanamo Munroe 
122. 4447. Euptoieta claudia (Cramer) 

123. 4449. Speyeria diana (Cramer) 

124. 4450. Speyeria cybele cybele (Fabricius) 

125. 4476a. Anthanassa texana seminole (Skinner) 
126. 4480. Phyciodes phaon (W. H. Edwards) 

127. 4481. Phyciodes tharos tharos (Drury) 

128. 4489. Charidryas gorgone gorgone (Hiibner) 
129. 4490. Charidryas nycteis nycteis (Doubleday) 
130. 4516a. Euphydryas phaeton ozarkae Masters 

181. 4522b. Basilarchia arthemis astyanax (Fabricius) 
132. 5423a. Basilarchia archippus floridensis (Strecker) 
133. 5423b. Basilarchia archippus watsoni dos Passos 
134. 4554. Anaea andria Scudder 

135. 4557. Asterocampa celtis (Boisduval and Leconte) 
136. 4562. Asterocampa alicia (W. H. Edwards) 

137. 4562.1. Asterocampa clyton (Boisduval and Leconte) 
138. 4563. Asterocampa flora (W. H. Edwards) 

139. 4568b. Enodia portlandia missarkae J. R. Heitzman and dos Passos 
140. 4568.1. Enodia anthedon anthedon A. H. Clark 
141. 4568.2 Enodia creola (Skinner) 

142. 4569. Satyrodes appalachia (R. Chermock) 

148. 4573. Cyllopsis gemma gemma (Hitbner) 

144. 4575. Hermeuptychia sosybius (Fabricius) 

145. 4576. Neonympha areolata areolata (J. E. Smith) 
146. 4578. Megisto cymela cymela (Cramer) 

147. 4587a. Cercyonis pegala abbotti F. M. Brown 
148. 4587b. Cercyonis pegala alope (Fabricius) 

149. 4614. Danaus plexippus (Linnaeus) 

150. 4615a. Danaus gilippus berenice (Cramer) 


LITERATURE CITED 


HopDGEs, RONALD W. (Editor). 1983. Check list of the Lepidoptera of America north 
of Mexico. E. W. Classey Limited and The Wedge Entomological Research Foun- 
dation, London. 284 pp. 

Howe, W. H. 1975. The butterflies of North America. Doubleday & Co. Inc., Garden 
City, N.Y. 633 pp. 

HUTCHINS, R. E. 1933. Annotated list of Mississippi Rhopalocera. Can. Entomol. 65: 
210-213. 

MATHER, B. & K. MATHER. 1958. The butterflies of Mississippi. Tulane Stud. Zool. 6: 
63-109. 

1959. The butterflies of Mississippi—Supplement No. 1. J. Lepid. Soc. 13:71- 





ize 





1976. The butterflies of Mississippi—Supplement No. 2. J. Lepid. Soc. 30:197- 
200. 

MILLER, L. D. & F. M. BRown. 1981. A catalogue/checklist of the butterflies of 
America north of Mexico. Lepid. Soc. Memoir No. 2. 280 pp. 

WEED, H. E. 1894. A preliminary list of the butterflies of northeastern Mississippi. 
Psyche 7:129-13]1. 


Journal of the Lepidopterists’ Society 
$9(2), 1985, 139-144 


A NEW SPECIES OF EXOTELEIA (GELECHIIDAE) 
REARED FROM PONDEROSA PINE 


RONALD W. HODGES 


Systematic Entomology Laboratory, USDA, % U.S. National Museum of Natural History, 
MRC-168, Washington, D.C. 20560 


ABSTRACT. Exoteleia anomala, new species, is described from New Mexico and 
Arizona. The larvae are needle miners on ponderosa pine. Problems with recognition of 
North American species of Exoteleia are discussed. 


A new species of Exoteleia was reared by R. E. Stevens from needles 
of ponderosa pine, Pinus ponderosa Douglas ex Lawson, near Silver 
City, Grant County, New Mexico. Reared adults were sent to me for 
identification. The moths proved to be an undescribed species that is 
most closely related to Exoteleia pinifoliella (Chambers). Exoteleia 
anomala Hodges is described to permit discussion of it and related 
species. 


Exoteleia anomala, new species 


A small dark-brown to black and pale-gray banded moth (Fig. 1). 
Most scales have shining yellowish reflections depending on angle of 
light incidence. 


Description. Head: haustellum white, several gray-tipped scales basally; labial palpus 
mainly white, lateral surface of first and second segments with many dark brown-tipped 
scales, inner surface of second segment with a few dark brown-tipped scales near apex, 
apex of second segment white, third segment with a partial ring of dark brown-tipped 
scales at 4 length and a well-developed ring of dark brown-tipped scales at % length; 
antenna, ventral surface mainly gray, scape off white ventrally and on anterior margin, 
dorsal surface dark brown, individual scales off white basally; shaft dark, alternate scale 
rows dark brown and gray; sensory cilia of male very short, scarcely visible at base of 
each segment at 100x magnification; frons, vertex, and occiput white, a narrow band of 
dark brown-tipped scales on anterior margin of eye, dark gray scales on posterior margin 
of eye. Foreleg: coxa and trochanter mottled pale and medium gray; femur darker gray; 
tibia dark gray, a few white scales at 4% length, % length, and apex; tarsus dark gray, base 
and apex of some scales paler; base and apex of Ist tarsomere with white scales, apex of 
2nd and 5th tarsomeres with off-white scales. Midleg: similar to foreleg, apex of each 
tarsomere with white scales. Hindleg: coxa and trochanter off white; femur mottled pale 
and dark gray; tibia mottled pale and dark gray, dorsal tuft of long scales off white, 
outer spurs mainly off white; tarsus mottled off white and dark gray, base and apex of 
lst tarsomere and apex of other tarsomeres white. Thorax mottled dark and pale gray- 
brown, individual scales with pale apexes and pale ridges. Wings: upper surface as 
illustrated; forewing mottled dark gray brown to black and pale gray to white; patches 
of upturned scales at approximately 4, % and % length; ventral surface of forewing with 
linear zone of dark brown scales (male only) that have the scale apexes directed toward 
the posterior margin, zone extending from before % length of wing nearly to posterior 
margin behind apex. Wing length: 4.9 mm (4.0-5.0 mm). Abdomen: dorsal and ventral 
surface of segments dark brown medially, off white laterally and distally. Male genitalia: 
as in Figs. 2 and 4. Female genitalia: as in Fig. 3. 

Types. Holotype: male, New Mexico, 40 km NE Silver City; Pinus ponderosa, vi.1977; 


140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Exoteleia anomala, new species, holotype male. 


R. Stevens; Hopkins U.S. #36961. Paratypes: 11 males, 8 females; same data as for 
holotype; USNM genitalia slides #10893-10902. 2 males, 4 females; Arizona, 10 km N 
Fort Apache; Pinus ponderosa, J. M. Schmid; Hopkins U.S. #66729, reared 8/82; USNM 
genitalia slides 11745-11748. In collection U.S. National Museum of Natural History. 

Host plant. Pinus ponderosa Douglas ex Lawson. 

Variation. The description is based on the holotype. Major variation occurs in the color 
of the transverse dark fasciae on the forewing that may be dark gray brown to shining 
red orange brown. Some specimens have gray-marked scales on the vertex and occiput. 


Discussion. Males of Exoteleia anomala can be recognized to genus 
by the series of dark brown raised scales on the under surface of the 
forewing. These scales are directed somewhat transversely with the 
long axis of the wing. Exoteleia anomala is nearest pinifoliella (Cham- 
bers) in genital characters; perhaps neither sex can be separated from 
pinifoliella consistently on them; males definitely cannot. The general 
coloration of the upper surface of the forewings and thorax of anomala 
is gray brown as viewed with the eye as contrasted with the warm red 
brown or brown of pinifoliella. Pinifoliella is known from southern 
Ontario and the New Jersey Pine Barrens, south along the Appalachian 
Mountains to Georgia, and from the Boston Mountains in northwestern 
Arkansas. Anomala occurs in New Mexico and Arizona. 

When specimens of anomala were sent to me for identification, I 
anticipated writing a key to adults of species of Exoteleia; however, I 
have been utterly frustrated in an attempt to do so. In addition to the 
introduced dodecella (Linnaeus), pinifoliella, burkei Keifer, chillcotti 
Freeman, and nepheos Freeman occur in North America. Exoteleia 


14] 


VOLUME 39, NUMBER 2 





Exoteleia anomala, genitalia: 2, 4, male; 3, female. 


Fics. 2-4. 


142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


graphicella (Busck) and californica (Busck) are not congeneric with 
dodecella and will be transferred at a future date. Dodecella is a large 
species (4.8-5.7 mm wing length) and is distinctly gray. It occurs in 
southern Ontario, Maine, and New York. Martin (1959) published on 
its bionomics for southern Ontario. 

Study of genitalia of 80 specimens from populations throughout the 
range of the native species has shown that none of the genital char- 
acters cited by previous authors is significant to discriminate among 
species. In males the margin of the lightly sclerotized part of the uncus, 
the size and shape of the mediolateral lobes from the saccus, the rel- 
ative length and shapes of the valvae and lobe from the posterior 
margin of the saccus all vary independently of other characters. In 
females the length of the extended external genitalia from the apex of 
the ovipositor to the anterior apex of the apophyses anteriores relative 
to the length of the first seven abdominal segments seems to allow for 
some grouping of entities. The genitalia cluster in groups from % to 
nearly equal to the length of the first seven segments of the abdomen. 
Pinifoliella and anomala have relatively long female genitalia, with 
pinifoliella having slightly the longer genitalia. Forewing coloration, 
host plants, and geographic distribution separate pinifoliella and 
anomala. What appears to be an undescribed species is small, dark, 
and has the shortest female genitalia relative to the first seven abdom- 
inal segments of the native species. It occurs in eastern North America 
from Lakehurst, New Jersey and Ithaca, New York, south to Mc- 
Clellanville, South Carolina, the southern Appalachian Mountains, and 
Hartford, Arkansas. The female genital group that includes nepheos 
has three very different looking moths: 1) the “large”? dark brown 
forewinged, dark gray-brown hindwinged nepheos; 2) an undescribed 
entity from the type series of pinifoliella (Ithaca, New York) that has 
relatively dark brown forewings and medium gray hindwings; and 8) 
a series of populations from South Carolina, Florida, and Louisiana 
that has relatively light orange-brown and off-white banded forewings 
and pale gray-brown hindwings. These populations are unlikely to 
represent one species. The fifth group includes burkei and what prob- 
ably is chillcotti from eastern Texas and Louisiana. The forewings of 
burkei are dark red brown, and the hindwings are dark gray brown; 
the forewings of the Texas specimens are pale orange brown and off- 
white banded, and the hindwings are very pale gray. 

On the basis of the material that I have studied I can defend and 
define four species, anomala, pinifoliella, dodecella, and an unde- 
scribed species from the eastern United States. I have been unable to 
define nepheos, burkei, chillcotti, and potentially two other entities on 
adult characters. 


VOLUME 39, NUMBER 2 143 


It would appear that pupal characters may be useful to define species; 
however, because voucher material is not available to support some 
published observations, I am unable to associate the differences noted 
in the literature with the moths that I have studied. The type series of 
pinifoliella contains three species: pinifoliella; the small, dark species; 
and one very much like pinifoliella but that is associated with nepheos 
by the female genitalia. Bennet (1966) illustrated the pupa of chillcotti, 
showing that it lacks the cutting plate of the pupa of what may be 
pinifoliella (Bennett, 1954). Because any of three very similar species 
probably occur in the Syracuse, New York area, it is not possible to 
state with certainty the species that he studied and called pinifoliella. 
This uncertainty points to the need for well-prepared voucher material 
to be deposited in permanent collections to document publications on 
life history studies of insects. Subsequent, finer or different, taxonomic 
conclusions could then be associated with previous literature. 

Larval behavior differs among the species. Burdick and Powell (1960) 
reported burkei as feeding on the needles of Pinus radiata D. Don. and 
P. sabiniana Dougl. into the fourth larval instar. Subsequently, the 
larva attacks the male staminate cones and rarely the developing buds. 
Stevens (1969), reporting on burkei (potentially) from Placerville, Cal- 
ifornia, indicated that the species fed on Pinus attenuata Lemm. In 
this infestation the last instar larvae attacked developing shoots and 
not staminate cones. Also, pupation occurred in the last larval habitat 
as contrasted with the larva usually leaving the last larval habitat to 
pupate as reported by Burdick and Powell (1960). Lindquist and Trin- 
nell (1967) found that the last instar larvae of nepheos fed on staminate 
cones and developing buds of Pinus resinosa Ait. and P. sylvestris L. 
and that pupation occurred in the last larval site. Freeman (1963) 
reported that chillcotti fed exclusively in needles of Pinus palustris 
Mill. and that pupation occurs there. Finnegan (1965) found the larva 
of pinifoliella feeding in the needles of Pinus banksiana Lamb.; and 
Bennett (1954) recorded P. rigida Mill., P. resinosa Ait., P. virginiana 
Mill., P. echinata Mill., P. palustris Mill., and P. pungens Lamb. as 
hosts. 

On the basis of available material I have been unable to resolve the 
question of separation of species in nearctic Exoteleia. I strongly urge 
that anyone who has the opportunity rear and preserve samples of the 
immature stages and adults of the local species. In New Jersey a light, 
larger species and a dark, smaller species are present. In the coastal 
plain of the Southeast two species may be sympatric. In Louisiana and 
Texas one, two, or three species occur. 

A generalized life history for all the species is that the moths appear 
to be univoltine with adults emerging from late spring to midsummer. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The adult female lays eggs in the entrance of an abandoned mine. 
Upon hatching, the larvae leave that site and attack other needles. 
Overwintering is in the larval stage, and the following spring the last 
instar larva may attack additional needles, staminate cones, or buds, 
apparently depending upon the species. Pupation usually occurs in the 
last instar larval feeding site; for burkei it may occur there or on or in 
the ground. 


ACKNOWLEDGMENTS 


I thank Molly K. Ryan for the line drawings that illustrate this paper, Douglas C. 
Ferguson for the photograph of the adult, and J. F. Gates Clarke and Raymond J. Gagné 
for review of the manuscript. 


LITERATURE CITED 


BENNETT, W. H. 1954. The pupal morphology of the pine needle miner (Lepidoptera: 
Gelechiidae). Proc. Entomol. Soc. Washington 56:41—42. 

1966. Pupal morphology of Exoteleia chillcotti Freeman (Lepidoptera, Gele- 
chiidae). Proc. Entomol. Soc. Washington 68:181-183. 

BURDICK, D. J. & J. A. POWELL. 1960. Studies on the early stages of two California 
moths which feed in the staminate cones of digger pine (Lepidoptera: Gelechiidae). 
Can. Entomol. 92:310-320. 

FINNEGAN, R. J. 1965. The pine needle miner, Exoteleia pinifoliella (Chamb.) (Lepi- 
doptera: Gelechiidae), in Quebec. Can. Entomol. 97:744—-750. 

FREEMAN, T. N. 1963. Two new species of coniferous needle miners from Louisiana 
and the description of a new genus (Lepidoptera: Gelechiidae). Can. Entomol. 95: 
727-730. 

LINDQUIST, O. H. & J. R. TRINNELL. 1967. The biology and description of immature 
stages of Exoteleia nepheos (Gelechiidae) on pine in Ontario. J. Lepid. Soc. 21: 
15-21. 

MARTIN, J. L. 1959. The bionomics of the pine bud moth, Exoteleia dodecella L. 
(Lepidoptera: Gelechiidae), in Ontario. Can. Entomol. 91:5-14. 

STEVENS, R. E. 1969. Occurrence of Exoteleia burkei in the Sierra Nevada (Lepidoptera: 
Gelechiidae). Pan-Pac. Entomol. 45:238. 





Journal of the Lepidopterists’ Society 
89(2), 1985, 145 


GENERAL NOTES 


A GENERIC REPLACEMENT NAME IN THE 
NACOPHORINI (GEOMETRIDAE) 


One of the twenty new generic names proposed in “A Generic Revision of the New 
World Nacophorini (Lepidoptera, Geometridae)” (Rindge, 1983, Bull. Amer. Mus. Nat. 
Hist. 175:147-262) was Azuayia. D. S. Fletcher of the British Museum (Natural History) 
was kind enough to remind me that this name had already been published by Dodge 
(1967, Pacific Insects 9:681). I hereby propose the replacement name Postazuayia for 
Azuayia Rindge, 1983, op. cit., p. 199; the type species remains Cidariophanes stigma- 
talis Dognin, and the gender is feminine. 


FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural 
History, Central Park West at 79th St., New York, New York 10024. 


Journal of the Lepidopterists’ Society 
39(2), 1985, 145-146 


THE CORRECT NAME FOR WHAT HAS BEEN CALLED 
LYCAEIDES ARGYROGNOMON IN NORTH AMERICA 


I have been asked by the Chairman of the Committee to Review/Update Memoir No. 
2 of the Lepidopterists’ Society to explain briefly the confusion that has arisen in North 
America over the name Lycaeides argyrognomon as used by various authors. I think the 
situation is sometimes misunderstood; so I shall review the circumstances in which the 
name has been used during the last 150 years and of the final identification of Berg- 
strdsser's butterfly, Lycaeides argyrognomon. Before doing so J must emphasize, to avoid 
confusion, the importance of using the correct, valid names for all animals. Rules for the 
formation and use of names have been the concern of zoologists at least since 1842, with 
rules and codes of practice usually agreed upon by committees of eminent specialists. 
The last International Code of Zoological Nomenclature was published in 1964 includ- 
ing Rules and Recommendations accepted by all countries. Its most important objectives 
are to maintain the Law of Priority and the Law of Homonymy. 

Turning now to the name argyrognomon, this was introduced into general butterfly 
nomenclature as Cupido argyrognomon in 1871 by W. F. Kirby in his Synonymic 
Catalogue of Diurnal Lepidoptera. From that time most European authors applied this 
name, as used by Kirby, to cover the European species of Lycaeides, before, of course, 
the presence of a second European species was discovered. In the early years of the 
present century the well-known French entomologist Charles Oberthiir became interest- 
ed in the group. The importance of genitalic structure in butterfly taxonomy was just 
becoming known, and Oberthiir realized the presence in Europe of two species of Ly- 
caeides, one of them a local insect, its male genitalia with long falces, which he named 
in 1910 as Lycaena argus ligurica. At the same time he wrote to two well-known spe- 
cialists, Prof. Courvoisier and Dr. Chapman in England, asking them to investigate 
“Lycaena argus and its forms, Races and Species.”’ Material for this investigation was 
obtained from many localities in Europe and Asia. The two entomologists selected by 
Oberthtir made independent reports which were included by Oberthiir in his private 
publication in 1917 (Lep. Comp. XIV:2-70), including a note from Dr. Reverdin about 
the genitalic structure in Lycaeides and numerous photographic plates. Both men agreed 
that two species were present in Europe. Prof. Courvoisier accepted Oberthiir’s name 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ligurica for the species with long falces so newly identified. Chapman described it as a 
“New European Lycaena” under the name argus sp. nov. 

As the years passed more names were published, difficulties arose, and the International 
Commission for Zoological Nomenclature (I.C.Z.N.) was asked to help with the situation 
reported by Courvoisier and Chapman. They were asked especially to select suitable 
names for those species recently identified by examination of the genitalia and to select 
a type species for the genus Lycaeides. All of this proved to be extremely difficult. It 
involved examination of a large number of specimens. In many cases no obvious scientific 
name existed, and finding one often called for considerable ingenuity; so, not surprisingly, 
the work took several decades. The first results were promulgated by the Commission in 
1945 (Opinion 169) and finally in 1954 (Opinion 269). Among the important decisions 
was the selection of the Linnean name idas 1761 for the commoner species of Lycaeides. 
This was made possible by suppressing the earlier appearance of the name for a species 
so badly described as to be impossible to interpret. In dealing with the second species, 
recently identified and with specific characters in the male genitalia, the Commission 
was fortunate when it was found and announced in Opinion 269 that these characters 
were actually present in specimens of Lycaeides from the Bruchkébler Wald in northern 
Germany as recorded by Bergstrasser in his original description. Lycaeides argyrogno- 
mon, therefore, must become the valid specific name for this species, with impressive 
priority to 1779, taking precedence before all others. 

During the long interval after the Commission began its work and the final Opinion 
in 1954, entomologists naturally continued to use the butterfly names with which they 
were familiar. It was during these years that V. Nabokov made many contributions, 
especially concerning the Lycaeides distributed in North America. 

When discussing what should now be called L. idas, he used the name “argyrogno- 
mon,’ not invalid for much of that time, but after the announcement of Opinion 269 
in 1954 the position changed radically. This name was restricted to a newly identified 
European butterfly. Applied to any other species (e.g., L. idas) would be to create a 
misidentification, and to use it so must be entirely against the Rules. The correct use of 
the name Lycaeides idas L. proposed by the Commission has been accepted everywhere 
in Europe and Asia. I cannot understand why it has not been accepted in North America. 

Catalogue headings for Europe and America should be as follows: 


Lycaeides argyrognomon Bergstrasser 1779 (Papilio). Europe. 
syn: ligurica Oberthtir 1910 (Lycaena); syn: aegus Chapman 1917 (Plebejus). 
Lycaeides idas Linnaeus 1761 (Papilio). Palaearctic Region & North America. 
syn: scudderi W. H. Edwards 1861 (Lycaena); syn: anna W. H. Edwards 1861 (Ly- 
caena); syn: argyrognomon sensu Kirby et Auctorum, nec Bergstrasser [misidentifi- 
cation]. 
Lycaeides melissa W. H. Edwards 1875 (Lycaena). North America, ?E. Siberia, ?Japan. 


LIONEL G. HIGGINS, Focklesbrook Farm, Chobham, Woking, Surrey, England GU24 
SHB. 


Journal of the Lepidopterists’ Society 
39(2), 1985, 146-150 


TIGRIDIA ASESTA (LINNAEUS) (NYMPHALIDAE) IS NOT ASSOCIATED 
WITH THEOBROMA CACAO L. (STERCULIACEAE) 


The medium-sized (spread wingspan, tip-to-tip, 4.5 cm) orange, brown and white 
nymphalid butterfly, Tigridia asesta (Linnaeus), is broadly distributed in moist-to-wet 
tropical forests of Central and South America (Seitz, 1904, Macrolepidoptera of the 
World, Vol. 5: The American Rhopalocera, A. Kernan, Stuttgart). A general description 


VOLUME 39, NUMBER 2 147 


of the larva has been given, and the larval food plant reported to be “cocoa (Theobroma)” 
in the Upper Amazon (Seitz, op. cit.). In this note I report that the larval food plant of 
this butterfly in eastern Costa Rica is Pourouma sp. (Moraceae) and that the larva will 
also feed successfully on Cecropia sp. in the same family. My description of the larval 
stages also differs from that reported in Seitz (op. cit.). Based upon these preliminary 
natural history observations in Costa Rica, I conclude tentatively that the larval food 
plant record in Seitz (op. cit.), the only known food plant record for this species, is 
erroneous. Based upon the leaf size and shape in moraceous plants, it is easy to understand 
the mistaken identity of the food plants as being Theobroma (Sterculiaceae). 

On 29 February 1984 two spinose, nymphalid-like larvae were discovered on a 0.5 m 
tall moraceous seedling along a weedy roadside at “Finca La Tirimbina,” near La Virgen 
(10°23’N, 84°07'W; 220 m elev.), Heredia Province. The locality is within the ““premon- 
tane tropical wet forest” zone of the Atlantic watershed and Sarapiqui District. The 
entire seedling was uprooted and placed in an airtight, clear-plastic bag. In this manner 
both larvae were reared, one to adulthood. The bottom of the bag was lined with a few 
layers of soft paper to absorb excess moisture and fecal material, and this paper was 
removed and replaced every 2-3 days. The larval “culture” was transported from this 
locality to other points in Costa Rica during the rearing period. The single adult obtained, 
along with its pupal shell and two larval head capsules, are deposited in the entomological 
collections of the Milwaukee Public Museum. During the rearing period, the one larva 
that eventually pupated was transferred from the original food plant to fresh meristems 
of Cecropia sp. (Moraceae) collected near Turrialba, Cartago Province, Costa Rica. 

The third, fourth, and fifth larval instars, and the pupa, of T. asesta are shown in Fig. 
1. Information on earlier life stages was not available. What follows is an overall com- 
posite description of larval and pupal stages based upon this material. Chaetotaxy is not 
given since I am not erecting a key to related species or genera. The larval descriptions 
are consistent with the format and depth of such information as presented on recent 
pages of this journal. 

Third instar. 10-18 mm long (Fig. 1) with glossy black head capsule. Head capsule 
with one pair of 4 mm long scoli curved slightly posteriorly. Each head scolus with 
extensive number of short black spines along entire length. Body coloration generally 
brownish orange and all segments with black spines. Spine distribution the same as to be 
described below for the fifth instar. The anterior edge of each trunk segment bears a 
pair of small white dots dorso-laterally. All feet black. 

Fourth instar. Very similar to third instar except body coloration now velvety black 
(Fig. 1). This instar attains a body length of about 25 mm prior to molting to the fifth 
(final instar). 

Fifth instar. Attains a final length of 40 mm before pupation. Head capsule glossy 
dark red and notably bilobed vertically into two halves. Each head scolus about 7 mm 
long, curved slightly posteriorly, and arising from the apex of each head capsule “lobe.” 
Head scolus with white tip and many black spines of varying lengths. Coloration of trunk 
region now a dull reddish brown, almost a pale maroon hue. Body with many small 
white spots, each ringed in black, and giving the trunk region a speckled appearance 
(Fig. 1). Head capsule width now 2.2 mm and entire structure adorned with many small 
black spines. First thoracic segment with dorsal raised rectangular black “plate” bearing 
one pair of setae curved anteriorly. Pair of latero-ventrally located white spots ringed in 
black and with tiny black seta arising near each. Posterior edge of segment with a few 
small white dots (“flecks”). Second thoracic segment with two pairs (one dorsal and one 
lateral) of long, branched black spines; each of these spines with four small branches or 
spinelets. Immediately anterior to the spines one pair of dorsally located white dots ringed 
in black; posteriorly with dorso-lateral doublet of these spots and with a very small white 
dot immediately behind each of the dorsal-most members of the two doublets. Additional 
ringed white spot located latero-ventrally on each side. Third thoracic segment identical 
to the second. All thoracic spines about 4 mm long and all with the four spinelets about 
¥, the way down from tips. 

First abdominal segment bears markings similar to thoracic segments 2 and 3, but now 
with three pairs of black spines, all the same length. Same arrangement of spines repeated 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 














Fic. 1. Clockwise, from upper left photograph: third, fourth, and fifth larval instars, 


and pupa, all in situ, for Tigridia asesta (Linnaeus) (Nymphalidae) from the Sarapiqui 
District of Costa Rica. 


on all remaining abdominal segments, except on the ninth in which white spots are 
absent and only one pair of reduced black spines is present. Anal plate is brownish and 
clasper black. The most conspicuous difference between the fifth instar and the two 
earlier instars is the presence of orderly arranged white spots on the trunk segments in 
the former. 


VOLUME 39, NUMBER 2 149 


Pupa. The pupa is 27 mm long by 6 mm wide (dorso-ventral axis) by 5 mm thick 
(laterally) and hangs from the edge of a partly eaten leaf of the food plant (Fig. 1). 
Overall, the pupa resembles an irregularly shaped wood chip slightly tinged with green 
meant to mimic moss. The head capsule area is adorned with one pair of 3 mm long 
“horns,” and most of the cuticle surface of the pupa has a rough texture. Head, thorax, 
and wing pad areas light brown; abdomen ventrally with a moss-like green color and 
anteriorly with an irregular blotch (somewhat oval in shape) of dark brown. Abdomen 
ventrally with three pairs of knob-like projections on posterior segments. Approximately 
half-way along the distal edge of each wing pad a small “flange” of cuticle directed 
outwards, both together resembling tiny paddles directed downward. Dorsally, thoracic 
area adorned with a pair of knob-like projections, dark brown in color. Virtually entire 
length dorsally along anterior—posterior axis shaded with dark brown resembling a thick 
stripe. Cremaster strongly flattened, concave in ventral perspective and light brown. Pupa 
readily makes quick, whip-like movements of abdominal area when touched. Eclosion 
takes place in 10 days under the rearing conditions employed here. In the single instance 
observed, the butterfly emerged at 1000 h, with wings fully expanded within 15 minutes. 

Larval behavior and food plant. The two larvae discovered on the Pourouma seedling 
(one early third instar and one fourth instar) occupied separate leaves. Both larvae rested 
on the ventral surfaces of leaves. Feeding occurred at the edge of a leaf, the larva 
remaining concealed from above on the ventral leaf surface. The larva does not construct 
silken pathways or nests on leaves. All of the six leaves of the food plant seedling appeared 
to be meristem or very fresh, and the two larvae were initially discovered on the largest 
leaves (each one about 19.5 x 7.5 cm, the latter for the greatest width). The larvae also 
readily accepted meristem leaves of Cecropia. With the roots kept moist in water-drenched 
paper towels and moss, the Pourouma seedling remained “‘lush”’ for almost three weeks 
in the plastic bag, facilitating the rearing of the larvae. 

Seitz (op. cit.) reports the larva (instar not mentioned) as being “light green, often 
tinged yellowish, with light green lateral stripe, beneath darker coloured, head and spines 
black; on cocoa (Theobroma).” He describes the pupa as being “greenish-yellow, red- 
toned with branched wing-like continuations on the head; small white points, green spikes 
and black markings.” My observations clearly differ from those of Seitz for T. asesta in 
Costa Rica, including the larval food plant. Meristem leaves of Pourouma bear a super- 
ficial resemblance to seedling leaves of Theobroma cacao L., particularly in terms of the 
pattern of venation and general oblongate shape of leaves. It is therefore not difficult to 
understand how the food plant could be misidentified without verification from a botanist 
knowledgeable about tropical vegetation. The leaves of an adult Pourouma tree take on 
a “stellate” appearance, markedly distinct from seedling leaves. The Seitz description of 
the larva and pupa presents a greater challenge. Three different alternatives are: (1) the 
descriptions are based upon newly molted individuals prior to cuticular-hardening; (2) a 
distinctly different subspecies or local variety of T. asesta is involved; and (8) a different 
species was being described. Based upon my limited data and the very limited amount 
of published information on T. asesta life cycle and natural history to date, it is not 
possible to determine which of the above alternatives is correct. I do conclude, however, 
that the Seitz food plant record is incorrect. As an evolutionary unit, the Moraceae are 
systematically far removed from the Sterculiaceae (e.g., Cronquist, 1981, An Integrated 
System of Classification of Flowering Plants, Columbia, New York, 1262 pp.). However, 
systematically unrelated groups of plants may have independently evolved chemical 
features rendering some members of each group to be acceptable as both oviposition 
sites and larval food plants for a particular species of butterfly (A. M. Young, unpub. 
data). 

While I did not test for larval feeding on older, mature leaves of the food plant or 
Cecropia, larvae readily accepted meristem or young leaves. However, Cecropia saplings 
in nature have the highest levels of herbivore damage to mature leaves (Coley, 1983, 
Ecology 64:426-433). Tigridia asesta might be a herbivore specialized for feeding on 
young leaves of Moraceae in tropical forests. Brown and Heineman (1972, Jamaica and 
Its Butterflies, Classey, London) point out that the accuracy of the systematic position of 
Tigridia (formerly Callizona, as in Seitz, op. cit. also) in the tribe Coloburini depends 


150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in part upon determination of natural history information, including larval food plant 
records. Species within this tribe such as Colobura dirce Linnaeus exploit moraceous 
plants such as Cecropia as larval food plants (Brown & Heineman, op. cit.). Thus, my 
record of T. asesta on Pourouma and its acceptance of Cecropia as well point to confir- 
mation of this genus within the Coloburini. A significant departure in the natural history 
between Tigridia and Colobura, however, is the clustered oviposition and larval gregar- 
iousness in the latter genus (Brown & Heineman, op. cit.) and the solitary early stages 
in the former as reported for the first time in this note. 

Susan Sullivan Borkin and Joan P. Jass discovered the larvae on the food plant, and 
Luis Poveda assisted with food plant determinations. 


ALLEN M. YouNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53238. 


ERRATUM 


In my recently published note appearing in this journal (J. Lepid. Soc. 38:237-242), 
Papilio birchalli in the three figure captions should be deleted and replaced with Papilio 
victorinus. During the preparation of revisions of this paper, I forgot to make these 
changes. 


Allen M. Young 


Date of Issue (Vol. 39, No. 2): 7 January 1986 





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CONTENTS 


MAINTAINING SPECIES INTEGRITY BETWEEN SYMPATRIC POPULA- 


TIONS OF HYALOPHORA CECROPIA AND HYALOPHORA 
COLUMBIA (SATURNIIDAE) IN CENTRAL MICHIGAN. James 
P. Tutile oe 


THE BIOLOGY AND IMMATURE STAGES OF SPHINGICAMPA AL- 
BOLINEATA AND S. MONTANA IN ARIZONA (SATURNIIDAE). 
Paul M. Tuskes 0 oo 


NEVADA BUTTERFLIES: PRELIMINARY CHECKLIST AND DISTRI- 


BUTION. George T. Austin 2... 
MITOURA MILLERORUM (CLENCH) AND ITS OCCURRENCE IN THE 
UNITED STATES (LYCAENIDAE). Kurt JORNSON ec 


TECHNIQUES FOR MAINTAINING A CULTURE OF THE BLACK 
SWALLOWTAIL BUTTERFLY, PAPILIO POLYXENES ASTERIUS 
STOLL (PAPILIONIDAE). Maureen Carter & Paul Feeny ..... 


THE BUTTERFLIES OF MISSISSIPPI—SUPPLEMENT No. 3. Bryant 
Mather & Katharine Mather... te 


A NEw SPECIES OF EXOTELEIA (GELECHIIDAE) REARED FROM 
PONDEROSA PINE, Ronald W. Hodges ...... ee 


GENERAL NOTES 


A generic replacement name in the Nacophorini (Geometridae). Frederick 
Hy Rend ge oe ee ee) i 


The correct name for what has been called Lycaeides argyrognomon in 
North America. - Lionel G. Higgins 00.) ee 
Tigridia asesta (Linnaeus) (Nymphalidae) is not associated with Theobroma 
cacao: \,. (Sterculiaceae). Allen.M. Young... 


FIRRATUM oe Fe AO NE 0 ar 


134 


139 


Volume 39 1985 Number 3 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





18 May 1986 


THE LEPIDOPTERISTS’ SOCIETY 


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Cover illustration: Micropylar end view (130) of the egg of Sericosema sp. (probably 
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JOURNAL OF 


Tue LeEpPIDOPTERISTS’ SOCIETY 


Volume 39 1985 Number 3 


Journal of the Lepidopterists’ Society 
39(3), 1985, 151-155 


A NEW SPECIES OF TILDENIA FROM ILLINOIS 
(GELECHIIDAE) 


RONALD W. HODGES 


Systematic Entomology Laboratory, USDA, % U.S. National Museum of Natural History, 
MRC 168, Washington, D.C. 20560 


ABSTRACT. Tildenia georgei Hodges, new species, is described from Illinois. The 
larvae are leaf miners on Physalis heterophylla var. ambigua. An illustrated identification 
key to adults of the four nearctic species of Tildenia is presented. 


A new species of Tildenia was discovered in southern Illinois by 
Paul Gross while he was conducting research on insects associated with 
Physalis heterophylla var. ambigua (Gray) Rydberg (Solanaceae). 
Reared adults were sent to me for identification. The moths proved to 
be an undescribed species that is closely related to Tildenia glochinella 
(Zeller) and Tildenia inconspicuella (Murtfeldt) and indistinguishable 
from them in maculation and other external features. Tildenia georgei 
Hodges is described to permit discussion of it and related species. 


Tildenia georgei, new species 


Description. A small, dark, gray-brown moth. Upper surface as in Fig. 1. Head: 
Haustellum with pale yellow scales basally; maxillary palpus pale yellow; labial palpus 
upturned, extending to vertex, scales of lateral surface dark gray, individual scales ter- 
minally margined with very pale gray, mesal surface pale yellowish gray, scales darker 
on third segment; third segment slightly shorter than second segment, apex acute; frons 
and vertex pale yellowish gray, scales with yellow and purple reflections, scales anterad 
of eyes dark gray before pale gray apexes, scales on occiput pale yellowish gray basally 
and apically, dark gray before apex; antennal shaft with alternate scale rows pale yellow 
and dark gray, apex pale yellow. Forewing: Length, 3.3-4.7 mm; upper surface mottled 
pale gray, dark gray, pale yellow, and pale yellowish orange, a row of yellowish-orange 
scales on fold from near base to % length of fold, fringe shades of gray; undersurface 
nearly uniformly dark gray on membrane, fringe pale yellowish gray at membrane, 
mottled gray elsewhere. Hindwing: Membrane evenly gray, fringe yellowish gray basally 
becoming darker beyond base; male without row of long scales from costal margin of 
dorsal surface. Foreleg: Scales on anterolateral surface nearly uniformly dark gray, tipped 
with very pale gray, apex of each tarsomere mainly pale gray. Midleg: Much as for 
foreleg, coxa paler, yellowish gray. Hindleg: Coxa mainly pale yellow with shining 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Tildenia georgei, new species, paratype male. 


reflections; femur pale yellow dorsally, becoming darker ventrally, all scales tipped with 
pale gray; tibia pale yellow basally and dorsally, mottled gray and pale yellow elsewhere, 
lateral spurs dark gray, scales narrowly margined with pale gray; tarsus mainly gray, 
apex of each tarsomere pale yellowish gray. Abdomen: Shining dark gray dorsally, distal 
row of scales on each segment paler than preceding scales; ventral surface pale yellowish 
gray medially, some mainly gray scales laterally. Male genitalia: As in Figs. 2, 4. Female 
genitalia: As in Fig. 7. Larva: Leaf miner on Physalis heterophylla var. ambigua (Gray) 
Rydberg. 

Types. Holotype: Male. Illinois, Mason County, Sand Ridge State Forest; collected 16 
August 1982, emerged 7 September 1982; leg. Paul Gross. Paratypes: 6 males, 6 females. 
Same data as for holotype (1 male, 1 female). Same locality and data as for holotype 
except collected on 3 September 1979 (5 males, 5 females). All specimens are in the 
collection of the U.S. National Museum of Natural History. 

Remarks. Tildenia georgei is very similar in appearance to inconspicuella, glochi- 
nella, and Keiferia lycopersicella (Walsingham). Males can be distinguished by their 
lack of a row of long scales on the costal margin of the dorsal surface of the hindwing. 
Distinguishing characters are in the genitalia of both sexes and are indicated in the key. 
The genitalia must be examined for specific and generic determination. 


Key to Adults of Nearctic Tildenia Species 


Ly) Males ich 0 ce Se Ne 2 
=~ Pemiales: suse nui en Nh el! ON a 
2. Apex of valva with long, slender, medially directed lobe (Fig. 

CPR opera Lo eg Reig a Nae a eR CM SL UMM ge altisolani (Keifer) 


— Apex of valva without such a lobe 
3. Saccus shorter than lateral width of vinculum, broadly rounded 
CEL) 0) ne satis 2 Cah cad LS doe glochinella 
— Saccus longer than lateral width of vinculum, lateral margins 
nearly parallel before narrowly rounded apex (Fig. 2) ............. 
4. Valva with apex slender, extending to narrowly acute tip (Fig. 
3); pair of annelar lobes posterolaterad of aedeagus small, 
lightly sclerotized: coe Mes niart OU UN Iai cere inconspicuella 
- Valva with apex broadly acute (Fig. 2); pair of annelar lobes 
posterolaterad of aedeagus well developed, prominent .... georgei 


VOLUME 39, NUMBER 3 153 





Fics. 2-6. Tildenia spp., male genitalia: 2, 4, T. georget; 3, T. inconspicuella, apex 
of left valva; 5, T. glochinella, saccus and ventral view of vinculum; 6, T. altisolani, 
apex of right valva. 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 






: ly 


Fics. 7-10. Tildenia spp., female genitalia: 7, T. georgei; 8, T. inconspicuella, apo- 
physes anteriores and base of bursa copulatrix; 9, T. glochinella, apophyses anteriores 
and base of bursa copulatrix; 10, T. altisolani, apophyses anteriores and base of bursa 
copulatrix. 


VOLUME 39, NUMBER 3 155 


5. Ductus bursae heavily sclerotized basally, projecting anteriorly 
as a cone or cylinder beyond the margin of apophyses anter- 
DIGS (CET Ge WS) etal a, lak Brees an lca lle Re Rn ee 6 
— Ductus bursae membranous basally (Fig. 9) 0. glochinella 
6. Ductus bursae heavily sclerotized basolaterally, sclerotized part 
projecting anteriorly less than % length of apophyses anter- 
NePRe Sm Ibioeayll\() meee tetera Man eine ee eee altisolani 
— Ductus bursae heavily sclerotized basally, sclerotized part pro- 
jecting anteriorly more than % length of apophyses anteriores 
IS. Tl) eee Oey oth us reais nila eet ap ea an a Zz 
7. Heavily sclerotized base of ductus bursae nearly cylindrical (Fig. 
(NN er er ee ee Tye inconspicuella 
— Heavily sclerotized base of ductus bursae conical, curved to the 
PenmraUeOrliya (kien), cote cen mae te tre eee, FO georgei 


Other species of Tildenia are known in North America. They too 
are superficially similar and to date can be distinguished by characters 
of the male and female genitalia only. Most of them are from Arizona, 
but this may reflect entomologists’ collecting, not the moths’ distribu- 
tion. I anticipate treating them in a subsequent fascicle of The Moths 
of America North of Mexico. 

This species is named for George Gross, father of Paul Gross. 


ACKNOWLEDGMENTS 
I thank Molly K. Ryan for the line drawings that illustrate this paper, Douglas C. 


Ferguson for the photograph of the adult and review of the manuscript, and J. F. Gates 
Clarke and W. W. Wirth for review of the manuscript. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 156-162 


OBSERVATIONS ON THE BIOLOGY OF 
PARNASSIUS CLODIUS (PAPILIONIDAE) IN THE 
PACIFIC NORTHWEST 


DavipD V. MCCORKLE 


Division of Math and Science, Western Oregon State College, 
Monmouth, Oregon 97361 


AND 


PAUL C. HAMMOND 
2435 E. Applegate, Philomath, Oregon 97370 


ABSTRACT. This paper examines the biology and life history of Parnassius clodius 
Menetries in the Pacific Northwest. Habitats used by the species include subalpine mead- 
ows high in the mountains and lowland rain-forests west of the Cascade Range. The 
primary larval foodplants belong to the genera Dicentra and Corydalis of the family 
Fumariaceae. Larvae in alpine habitats often display a gray-brown camouflage pattern 
that blends with the rocks of the habitat. However, larvae in lowland rain-forests display 
a conspicuous black and yellow-spotted pattern that appears to mimic the warning colors 
of polydesmid millipedes. Larval development in lowland habitats is completed within 
a single year, and pupation takes place inside a strong, well-formed silken cocoon. Male 
butterflies display a “rape” type of mating, with no evidence of courtship behavior or 
sexual pheromones. Tough, tear-resistant wings and a large female sphragis may be 
related to this sexual behavior. 


Parnassius clodius Menetries belongs to a genus that is considered 
to be relatively primitive within the Papilionidae (Tyler, 1975). These 
are the only butterflies that have a moth-like pupa enclosed within a 
silken cocoon. Because of the putatively “primitive” nature of these 
butterflies, their life history and ecology is of considerable interest. Of 
the three species of Parnassius found in North America, only P. clodius 
is uniquely endemic to this continent and is widely distributed in the 
western mountains from southern Alaska to central California, western 
Wyoming, and northern Utah (Ferris, 1976). Some details of the life 
history and ecology of this species are outlined by Edwards (1885), 
Tyler (1975), and Dornfeld (1980). During the past twenty years, the 
present authors have studied various aspects of P. clodius biology in 
Oregon, Washington, and western Wyoming, resulting in much addi- 
tional information. 


Ecology and Life History 


In terms of ecology, P. clodius occupies two distinctly different types 
of habitat. One consists of open subalpine meadows and rocky slopes 
above timberline at high elevations in the mountains. We have ob- 
served the species in subalpine meadows throughout western Oregon 
and Washington, and in Yellowstone National Park of Wyoming. We 


VOLUME 39, NUMBER 3 157i 


also observed the species on alpine talus slopes above timberline at 
Harts Pass, Okanogan County, Washington. 

However, the most frequent habitat of P. clodius in the Pacific 
Northwest is the lowland rain-forests extending from the western slope 
of the Cascade Range west to the Pacific Ocean. Although typically 
found in moist riparian habitats along forest streams and mountain 
valleys, the species was formerly found in the Portland and Seattle 
metropolitan areas and is still quite abundant in the low foothills sur- 
rounding the Willamette Valley in Oregon and the Puget Sound trough 
in Washington. This forest habitat extends from the 4000 ft. (1200 m) 
elevation down to sea level near the ocean. 

The primary larval foodplant in these coastal rain-forests is the wild 
bleeding heart Dicentra formosa Andr., which is very abundant in 
moist forest habitats along the West Coast. A second probable food- 
plant is Corydalis scouleri Hook., a relatively uncommon species. We 
have not yet observed P. clodius larvae on this plant in the field, but 
they accept it readily in the laboratory. At high elevations in the alpine 
habitat and east of the Cascades, Dicentra uniflora Kell. is a likely 
foodplant. This species is a known foodplant of P. clodius in northern 
California (John F. Emmel, pers. comm.). All of these plants belong to 
the family Fumariaceae, and it is probable that related species such as 
Dicentra cucullaria L. and Corydalis aurea Willd. would also provide 
acceptable foodplants. 

The female butterflies oviposit on and near the Dicentra plants. 
However, we have also observed females ovipositing on shrubs up to 
four feet above the Dicentra beds. Evidently a specific chemical em- 
anating from the foodplant is sufficient to induce oviposition anywhere 
in the general vicinity of the foodplant. The larvae develop within the 
egg shell but do not emerge from the egg until the following spring. 
Eggs deposited on shrubs usually reach the Dicentra beds when the 
shrubs drop their leaves in the fall. Foodplant records such as Viola 
and Rubus mentioned by Ackery (1975) are almost certainly in error 
and may be due to this indiscriminate oviposition by the females. 

Early instar larvae have small tubercles, but later instars are mostly 
smooth with fine hairs. The larvae stay hidden in debris at the base of 
the foodplant most of the time. Feeding takes place very rapidly, so 
the larvae are exposed from cover only briefly. Nevertheless, P. clodius 
is frequently parasitized by tachinid flies in many localities. Osmeteria 
are poorly developed in Parnassius larvae and are not as important 
for defense against predators compared to Papilio larvae. 

Parnassius clodius larvae display two very distinct color morphs. 
One form is black with a lateral row of bright yellow spots on each 
side of the body (Fig. 1). The form of these spots is highly variable, 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 








mvs: see. A : 
Fics. 1-3. Left (1), larva of P. clodius, black form, Benton Co., Ore. Middle (2), 


larva of P. phoebus, Yakima Co., Wash. Right (3), larva of P. clodius, gray-brown form, 
Castle Lake, Siskiyou Co., Calif. 


d 





Fics. 4-6. Left (4), Harpaphe haydeniana, Polk Co., Ore. Middle (5), open net 
cocoon and pupa of P. phoebus (behind thick Sedum stems in lower center). Right (6), 
well-formed cocoon of P. clodius cut open to reveal pupa ready to eclose. 


ranging from large round spots to long slender bars, or may be divided 
into several smaller spots. This color pattern is very similar to that of 
P. phoebus Fabr. (Fig. 2) and the Eurasian P. apollo L. (illustrated by 
Stanek, 1969). However, P. phoebus differs in having a second, more 
dorsal row of yellow spots on each side of the body. The second color 
form in P. clodius is gray-brown or pinkish gray with creamy yellow 
lateral spots and dorsal rows of narrow chevron markings equivalent 
to the dorsal row of spots seen in P. phoebus (Fig. 3). In our experience, 


VOLUME 39, NUMBER 3 159 


TABLE 1. Sequence of experiment testing the mimicry-model system of Parnassius 
clodius larvae and the millipede Harpaphe haydeniana as protection against the grass- 
hopper mouse Onychomys leucogaster. 


. Clodius larvae given to mouse—larvae eaten. 

. Millipedes given to mouse—millipedes bitten, producing defense odor detectable to 
observer, mouse then rejected millipedes. 

Meal worms given to mouse—worms eaten. 

Clodius larvae given to mouse—larvae sniffed and rejected. 

Adult meal worm beetles given to mouse—beetles eaten. 

Clodius larvae given to mouse—larvae sniffed, handled, finally eaten after long delay. 
Millipedes given to mouse—millipedes sniffed and rejected. 

Meal worms given to mouse—worms eaten. 


bo ee 


GID Ut oo 


the gray-brown form is dominant in alpine populations of P. clodius, 
for example at Harts Pass in Okanogan County, Washington and at 
Donner Pass in Nevada County, California. This morph appears to be 
a camouflage pattern that blends with the rocks in the alpine habitat. 
By sharp contrast, the black and yellow-spotted form is very conspic- 
uous, is dominant in the lowland rain-forest populations of P. clodius, 
and appears to mimic the warning colors of polydesmid millipedes 
such as Harpaphe haydeniana Wood (Fig. 4). These millipedes are 
very abundant in the moist, riparian habitats used by P. clodius larvae. 

Some populations of P. clodius are polymorphic for both larval color 
forms. For example, larvae sent to us by John F. Emmel from Castle 
Lake in Siskiyou County, California displayed both color forms. Like- 
wise, an adult female butterfly collected at Chinook Pass near Mt. 
Rainier National Park, Washington produced ten larvae, five of the 
black form and five of the pinkish gray form. In these, the black larvae 
retained the narrow yellowish dorsal chevrons of the gray larvae, a 
trait absent in most lowland black larvae. This ratio between the black 
and gray forms is suggestive of a simple Mendelian inheritance for 
these color morphs. However, the chevron markings are apparently 
controlled by a separate set of gene loci. 

In 1973, one of the present authors (McCorkle) conducted an ex- 
periment to test the predator protection of the mimicry-model system 
that apparently exists between lowland P. clodius larvae and the mil- 
lipede Harpaphe haydeniana. Grasshopper mice (Onychomys leuco- 
gaster Max.) from eastern Oregon were used as predators in this ex- 
periment, since these insectivorous rodents do not occur within the 
ranges of the butterfly or millipede and would have no prior experience 
with these arthropods. The sequence of this experiment is shown in 
Table 1. 

This experiment appears to demonstrate that the mimicry color pat- 
tern of lowland P. clodius larvae can give them a degree of protection 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


against predators, although predators may with sufficient experience 
learn to distinguish the larvae from millipedes. In nature, however, the 
millipedes are commonly exposed in the open, while P. clodius larvae 
are usually hidden and only briefly exposed during feeding. Thus, the 
mimicry may work quite well in nature, since predators would be 
expected to have abundant experience with the millipedes and little 
experience with the larvae. 

In lowland populations of P. clodius, development is completed in 
a single year. The larvae emerge from the egg shells during March 
and start to feed on the young Dicentra plants. Full larval development 
is reached usually by late April or May, followed by pupal develop- 
ment of several weeks, and adult butterfly emergence in June and July 
depending upon elevation. The pupa is short and rounded, dark brown 
in color, and quite similar to a saturniid moth pupa. It is enclosed 
within a strong, well-formed silken cocoon (Fig. 6). By contrast, the 
cocoon of P. phoebus is very loose and poorly formed (Fig. 5). Wilson 
in Ehrlich and Ehrlich (1961) has suggested that this terrestrial cocoon 
may be an adaptation to the harsh, alpine climate, rather than a prim- 
itive trait. However, the above observations do not support this idea, 
since the lowland P. clodius has the best formed cocoon, and the more 
alpine P. phoebus has a poorly formed cocoon. 


Sexual Behavior 


The mating system of P. clodius adults is quite interesting, since 
these butterflies display virtually no evidence of courtship behavior. 
Indeed, the males display a “‘rape”’ type of mating in which the males 
engage the females and copulate by brute force. In dramatic contrast, 
most higher butterflies display elaborate courtship rituals in mating, 
often involving specialized sexual pheromones in both the male and 
female. For example, Brower, Brower and Cranston (1965) have out- 
lined in detail the courtship patterns of Danaus gilippus Cramer. Like- 
wise, Speyeria butterflies display a very elaborate courtship ritual in 
which the males flutter around the females, stimulating the females 
with a sweet, musky smelling pheromone that is easily detected by the 
human observer. In turn, the females release a second pheromone that 
stimulates the male to twist his abdomen towards the female for actual 
copulation. There is evidence that the female pheromone of Speyeria 
is often species specific and frequently serves as a reproductive isolat- 
ing mechanism that prevents interspecific hybridizations (see Grey, 
Moeck & Evans, 1963). 

However, P. clodius males, upon sighting a female, chase rapidly 
after her and literally attack her from behind. Upon grasping the fe- 
male in mid-air, the male and female drop abruptly to the ground. 


VOLUME 39, NUMBER 3 161 


The female then lies limply on the ground, often with wings crumpled 
in the vegetation, while the male sits on top of her in copulation. 
Parnassius have extremely tough, tear-resistant wings, which may be 
an adaptation to this rough mating behavior. Otherwise, few females 
would survive mating with intact wings. Both of the present authors 
have independently observed this mating behavior in the field on sev- 
eral different occasions. Two additional anecdotal observations of mat- 
ing behavior may also be mentioned here. On one occasion in the field, 
McCorkle captured a virgin female in the first swing of the net, fol- 
lowed shortly by capture of a male in a second swing. Before the two 
butterflies could be removed from the net, they were already in cop- 
ulation. On a second occasion, a reared male was placed near a virgin 
female in a laboratory window. Upon seeing the female, the male 
immediately attacked and engaged her in copulation. We would sug- 
gest that the “rape” mating behavior observed in these butterflies may 
be a primitive trait, compared to the elaborate “courtship” mating 
behavior observed in most other groups of butterflies. 

Moreover, females of many so-called “primitive” butterflies carry 
an external sphragis or internal genital plug following mating in order 
to prevent subsequent matings by additional males. Mated females of 
P. clodius carry one of the largest and best developed sphragis struc- 
tures seen in butterflies. By contrast, the sphragis of P. phoebus is much 
smaller (see illustrations in Tyler, 1975 and Dornfeld, 1980). Presence 
of a sphragis may be typical of butterflies with a “rape” type of mating 
behavior, since older, mated females are often resistant or non-respon- 
sive to males in species with a “courtship” mating type. 

Scott (1973) has made similar observations of the mating behavior 
in P. phoebus. He has suggested that virgin females of Parnassius may 
release a sexual pheromone attractive to the males, and that the females 
cease to produce this pheromone after mating and attachment of the 
sphragis. As a consequence, the males do not waste time and energy 
pursuing mated females. This possible pheromone system may well 
exist in P. clodius, but further studies are presently needed for confir- 
mation. Such a pheromone system could be species specific and serve 
to prevent interspecific hybridization between P. clodius and P. phoe- 
bus when sympatric. However, females of P. clodius also differ from 
those of P. phoebus in having the dorsum of the abdomen completely 
naked, which may also be important for species recognition during 
mating. 

In terms of relationships, it is quite possible that P. clodius represents 
a relatively generalized, primitive condition within the genus, consid- 
ering the lowland habitat, well-formed cocoon, and large female sphra- 
gis. Parnassius clodius is closely related to a group of Eurasian Coryda- 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lis-Dicentra feeders which include P. eversmanni Menetries, P. 
mnemosyne L., P. stubbendorfti Menetries, and P. glacialis Butler. It 
is interesting to note that these last two species are also reported to 
feed upon Aristolochia (Ackery, 1975), the same foodplant used by 
such related genera as Archon and Parnalius (=Zerynthia). In contrast, 
the Sedum-Saxifraga feeders such as P. phoebus, P. apollo, P. bremeri 
Bremer, and P. nomion Waldheim appear to be more specialized in 
habitat, foodplant, cocoon development, and female sphragis. 


ACKNOWLEDGMENT 


We would like to thank John F. Emmel for sending us samples of P. clodius larvae 
from California and for stimulating our interest in these early stages. 


LITERATURE CITED 


ACKERY, P. R. 1975. A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bull. Br. Mus. Nat. Hist. 31: 73-105. 

BROWER, L. P., J. V. Z. BROWER & F. P. CRANSTON. 1965. Courtship behavior of the 
queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 50:1-39. 

DORNFELD, E. J. 1980. The butterflies of Oregon. Timber Press, Forest Grove, Oregon. 
276 pp. 

EpDwarps, W. H. 1885. Description of some of the preparatory stages of Parnassius 
smintheus Doubl. and of P. clodius Men. Can. Entomol. 17:61-65. 

EHRLICH, P. R. & A. H. EHRLICH. 1961. How to know the butterflies. Wm. C. Brown 
Co., Dubuque, Iowa. 262 pp. 

Ferris, C. D. 1976. A note on the subspecies of Parnassius clodius Menetries found 
in the Rocky Mountains of the United States (Papilionidae). J. Res. Lepid. 15:65-74. 

Grey, L. P., A. H. MoECK & W. H. Evans. 1963. Notes on overlapping subspecies. II. 
Segregation in the Speyeria atlantis of the Black Hills (Nymphalidae). J. Lepid. Soc. 
17:129-147. 

ScoTT, J. A. 1973. Mating of butterflies. J. Res. Lepid. 11:99-127. 

STANEK, V. J. 1969. The pictorial encyclopedia of insects. Paul Hamlyn, London. 544 


Pp. 
TYLER, H. A. 1975. The swallowtail butterflies of North America. Naturegraph Pub- 
lishers, Inc., Healdsburg, California. 192 pp. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 163-170 


THE BIOLOGY AND IMMATURE STAGES OF 
AUTOMERIS RANDA AND AUTOMERIS IRIS HESSELORUM 
(SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge 111D, Houston, Texas 77054 


ABSTRACT. The biology and immature stages of Automeris randa and Automeris 
iris hesselorum are described for the first time. Mature larvae and adults of both species 
are illustrated and a key to the Automeris larvae of the United States is presented. 
Automeris randa occurs in the Pelincillo Mts. of Arizona and New Mexico. The larval 
food plants of A. randa include Quercus and Celtis. Automeris iris hesselorum has been 
collected near Pena Blanca Lake, Santa Cruz Co., Arizona. The larval food plants of A. 
iris hesselorum include Quercus and Eysenhardtia polystachya. Both species are single 
brooded in Arizona. 


Although Automeris is a large neotropical genus, few members oc- 
cur north of Mexico. To date, only six species of Automeris are known 
to occur in the United States and only one species, Automeris io (F.) 
is widely distributed. Automeris io occurs from southern Canada to 
Mexico and inhabits most of the contiguous states with the exception 
of those in the far west. Automeris louisiana Ferguson & Brou and A. 
zephyria Grote are endemic species, while A. cecrops pamina (Neu- 
mogen), A. randa Druce, and A. iris hesselorum Ferguson are pri- 
marily Mexican species that reach their northern limit of distribution 
in Arizona or New Mexico. Ferguson (1972) noted that the biology and 
immature stages of A. iris hesselorum were undescribed, and the oc- 
currence of A. randa in the United States was not known until 1976, 
thus, this species was not illustrated or discussed by Ferguson. The 
purpose of this paper is to describe the biology and immature stages 
of A. randa and A. iris hesselorum. 


Automeris randa 


Automeris randa is widely distributed in Sonora and Chihuahua, 
Mexico, but in the United States is known only from the area near the 
Arizona—New Mexico—Mexico border. The first United States record 
was that of a male captured by Peter Jump in 1976 at Guadalupe 
Canyon, Cochise Co., in the Peloncillo Mts. of Arizona. Between 1976 
and 1981 numerous trips were made to the area, but only a few ad- 
ditional males were captured. But, during this time, randa was also 
collected at Cottonwood Canyon, 1440 to 1600 m, Cochise Co., Arizona 
and Clanton Draw, 1660 m, Hidalgo Co., New Mexico by Jump. Both 
of these locations are also in the Peloncillo Mts. Then in 1982 and 1983 
both males and females were captured by numerous collectors at Gua- 


164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


dalupe Canyon, and a specimen was taken in the same mountain range 
at Skeleton Canyon. 

The adults (Fig. 2) have apricot colored forewings, while the mar- 
ginal area is light brown. Some females have a slight purplish tint on 
the forewing. Two thin lines, yellow and brown, extend from near the 
apex of the forewing to the inner margin. The thin antemedial line is 
yellow and the base of the wing is dusted with white scales. On the 
hindwing, the margin is light brown and there is a reddish brown to 
orange submarginal band that is distal to a thin black and thin yellow 
line. The basal and medial hindwing area is orange, and surrounds a 
large multi-colored eyespot. The thorax is dark brown, and the abdo- 
men is orange. Forewing length of the males ranges from 42 to 48 
mm; females from 44 to 55 mm. According to LeMaire (pers. comm.), 
randa may represent a subspecies of A. rubescens Walker, but a change 
is not contemplated unless additional work is done on this complex. 

Capture records from Arizona and New Mexico indicate that adults 
have been taken at lights between 30 June and 7 August. Based on 
these records, the peak flight appears to be from late July through 
early August. In captivity, adults emerged between 0900 and 1500 h. 
After wing expansion they remain quiescent throughout the day. Mat- 
ing generally occurs after 2200 h and the pair remain together for 
about 45 min. The female begins the ovipositional flight the next night. 
The white eggs are deposited in clusters of 20 to 45 on the underside 
of the leaf. 

The larvae feed gregariously until the 4th or 5th instar and then 
singly or occasionally in pairs. Early instar larvae are yellow but be- 
come green or blue-green at the onset of the 4th or 5th instar and 
develop a prominent yellow subspiracular line that is usually bordered 
on either side by a thin black and red line. The dorsal and dorsolateral 
scoli are relatively long and slant forward during the 5th—7th instars, 
giving the larva a distinct appearance (Fig. 1). Although little pheno- 
typic variation was observed, it was found that the number of larval 
instars varied from 6 to 7. The larvae having the additional instar had 
larger head capsules and greater overall size and, thus, may have been 
females. First through last instar larvae collected in Guadalupe Canyon 
have been found on various oaks and hackberry, Celtis pallida Torr. 

Prior to pupation, larval coloration may change from green to yellow 
or yellowish brown. In captivity all larvae pupated on the host plant 
by attaching leaves together and spinning a thin but tightly woven 
cocoon. Neither the cocoon nor the leaves are attached to the branch 
with silk and would have fallen into the leaf litter when the tree shed 
its leaves. 

In addition to A. randa, other species of saturniids collected during 


VOLUME 39, NUMBER 3 165 





Fics. 1-4. 1, mature larva of A. randa (0.65 x ); 2, male A. randa (0.65 x ); 3, mature 
larva of A. iris hesselorum (0.85 x); 4, male A. hesselorum (1.0x). 


late July at Guadalupe Canyon included: Hyalophora gloveri, Eupack- 
ardi calleta, Antheraea polyphemus oculea, Agapema galbina anona, 
Citheronia splendens, Anisota oslari, Sphingicampa hubbardi, and 
Automeris cecrops pamina. 

The larval description is based on 26 larvae reared from ova depos- 
ited by a female collected at Guadalupe in 1983 and on four second 
instar field collected larvae provided by Steve McElfresh in 1982 and 
reared to maturity by the author. Length and width measurements 
give the size range of larvae at the end of each instar. 


Larval Description 


First instar. Head: Diameter 1.0 mm. Color yellow. Body: Ground color yellow. 
Length 5.6 mm, width 1.0 mm. Dorsal scoli branched, shaft and spines yellow w ith black 
tips. Dorsolateral, lateral and sublateral scoli yellow. Dorsal, lateral and ventral surfaces 
yellow. Thin mid-dorsal brown line extends from mesothorax to caudal segment. True 
legs and prolegs yellow. 

Second instar. Head: Diameter 1.7 mm. Color yellow. Body: Ground color yellow. 
Length 10 mm, width 1.6 mm. Dorsal scoli yellow, spines with black tips. Dorsolateral, 
lateral and sublateral scoli yellow. Two thin black lines extend length of larva. Mid- 
dorsal line extends from metathoracic segment (T3) to last abdominal segment (AQ). 
Lateral line mid-way between dorsolateral and lateral scoli extending from T3 to AQ; 
interrupted by yellowish brown intersegmental area. Dorsal, lateral and ventral surfaces 
yellow. True legs and prolegs yellow. 

Third instar. Head: Diameter 2.4 mm. Color yellow. Body: Ground color yellow. 
Length 13-15 mm, width 2.1 mm. Dorsal and dorsolateral scoli branched, shaft and 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


spines yellow, some with black tips. Lateral and sublateral scoli branched and yellow. 
Three thin black lines extend length of larva. Mid-dorsal and dorsolateral lines as in 
second instar. Black spiracular line extends from T1 to A9. Dorsal, lateral and ventral 
surfaces yellow. Spiracles black. True legs and prolegs yellow. 

Fourth instar. Head: Diameter 3.0 mm. Color yellowish green. Clypeus light brown. 
Body: Ground color yellowish green. Length 19-21 mm, width 4.5 mm. Dorsal and 
dorsolateral scoli yellow, tip of shaft and spines black. Lateral and sublateral scoli branched, 
shafts and spines yellowish green. Dorsal and lateral surfaces with numerous thin lines 
extending length of larva. Line I, mid-dorsal black line bordered on either side by thin 
yellow line. Line II, greenish gray line passes just below base of each dorsolateral scoli. 
Line III-IV, two lines, yellow above black, pass between dorsolateral and lateral scoli. 
Dorsal projection from black line disrupts yellow line as it pass each scoli. Line V, gray- 
green line connects base of each lateral scoli. Line VI, yellow line passes just ventral to 
line V and lateral scoli. Ventral surface gray-green. Spiracles black. Prolegs light yellow 
with light brown shields. True legs gray-green. 

Fifth instar. Head: Diameter 4.1—-4.7 mm. Color green to blue-green with short white 
setae. Clypeus cream. Body: Ground color green to blue-green. Length 27-34 mm, width 
8-9 mm. All scoli shafts and spines match ground color. Dorsal and dorsolateral thoracic 
scoli shafts with upper % black. Dorsal abdominal scoli with black tips on some spines. 
Lateral and sublateral scoli reduced in size. Lighth bluish gray mid-dorsal line extending 
length of larva and edged by thin light yellow on either side. Thin short black line 
extends from base of dorsolateral scoli to dorsal scoli and then between dorsal scoli. Black 
spiracular line thin and disrupted by base of spiracle, subspiracular yellow and black 
spiracular lines extend from T3 to AQ. Small red patch with short white setae extending 
from cream colored pinaculum in intersegmental area of sublateral surface just anterior 
of each proleg and on A7. Anal and proleg shields light brown to light orange with 
numerous short white setae extending from cream colored pinaculum. Prolegs and ven- 
tral surface green. Spiracles light brown. True legs light orange. 

Sixth instar (Fig. 1). Head: Diameter 5.5-5.9 mm. Color green to blue-green with 
short white setae. Clypeus cream. Body: Ground color green or blue-gray. Length 52- 
56 mm, width 12 mm. Dorsal and dorsolateral scoli enlarged (7 mm) with thin shafts 
and numerous long thin spines (4 mm). Lateral and sublateral scoli reduced (2-2.5 mm) 
in size. All scoli match ground color; no black tipped spines present as in previous instar. 
Short black vertical bar occurs between the dorsal and dorsolateral scoli. Broad yellow 
subspiracular line below thin black spiracular line which is disrupted by lower edge of 
spiracle. Ventral edge of subspiracular line often thinly edged with black and red. Both 
lines extend from anterior edge of T3 to A9. Red patch with short white setae extending 
from cream colored pinaculum in intersegmental area of sublateral surface just anterior 
to each proleg and A7. Anal and proleg shields reddish brown with white setae extending 
from cream colored pinaculum. Spiracles light orange. True legs brownish orange. 

Seventh instar. Head: 7.9-8.1 mm. Color and comments same as previous instar. Body: 
Length 72-78 mm, width 17-19 mm. Ground color green or blue-gray. Dorsal and 
dorsolateral scoli; shafts 9-11 mm, spines to 6 mm. There is no noticeable difference 
between sixth and seventh instar larvae, other than overall size. 


Automeris iris hesselorum 


Like the previous species, Automeris iris hesselorum has an ex- 
tremely limited distribution in the United States. Ferguson (1972) il- 
lustrated the adults in color and indicated that all of the Arizona rec- 
ords were from Pena Blanca Lake, Santa Cruz Co. This species was 
taken with some frequency until 1969, but available records suggest 
that none have been captured at Pena Blanca Lake since 1972. In 
addition to Pena Blanca Lake, a single specimen has been taken at San 
Rafael Valley, Cochise Co., Arizona. The dates of capture range from 


VOLUME 39, NUMBER 3 167 


mid July to early August. Similarly, a large series taken in Temoris, 
Sonora, Mexico were collected between 19 July and 8 August. The 
flight period probably extends for a week or two on either end of the 
known season. 

The adults are easy to separate from the other two species of Au- 
tomeris occurring in Arizona. Automeris iris hesselorum has a wing 
shape similar to that of Automeris io. The forewings are brownish 
pink, and the dark brown postmedial line extends from the costa to 
the inner margin in a straight line and remains the same distance from 
the wing margin (Fig. 4). The wing span of the male varies from 57 
to 65 mm; female from 68 to 74 mm. The forewings of A. randa and 
pamina are falcate or pointed and the yellowish medial line extends 
diagonally from near the apex to about midpoint on the inner margin 
of the wing. 

It would appear that the larvae of hesselorum utilize a wide variety 
of food plants. Ron Wielgus (pers. comm.) collected a late instar hes- 
selorum larvae on an unidentified species of oak at Pena Blance Lake 
in 1968. During this same period, larvae were reared in captivity on 
various Quercus species, and desert willow, Chilopsis linearis (Cav.) 
which is in the family Bignoniaceae and not related to willow as the 
common name suggests. Larvae of hesselorum were collected in Mex- 
ico on Kidneywood, Eysenhardtia polystachya (Ort.) but when trans- 
ferred to oak failed to survive (Steve Prchal, pers. comm.). 

First through 4th instar larvae are a uniform golden yellow and their 
bodies are slightly tapered at each end. Early instar larvae feed gre- 
gariously on the underside of the leaves. Fifth and 6th instar larvae 
are green and appear somewhat similar to larvae of Automeris io. 
Mature larvae have a prominent white subspiracular line below thin 
black and red lines that extend from the first abdominal segment to 
the caudal segment (Fig. 3). The larval description is based on three 
larvae reared to maturity by the author on oak. The three fertile ova 
were received from Steve Prchal, who had collected a female iris hes- 
selorum in Sonora, Mexico. 


Larval Description 


First instar. Head: Diameter 0.9 mm. Color golden yellow. Body: Ground color golden 
yellow. Length 4.9 mm, width 1.2 mm. Scoli coloration matches ground color. 

Second instar. Head: Diameter 1.7 mm. Color golden yellow. Body: Ground color 
golden yellow. Length 11 mm, width 2.1 mm. Scoli coloration matches ground color. 

Third instar. Head: Diameter 2.5 mm, color golden yellow. Body: Ground color 
golden yellow. Length 21-24 mm, width 4.5 mm. All scoli with golden yellow spines 
and shafts. Dorsal and dorsolateral scoli with some spines tipped with black. Light blue- 
gray mid-dorsal line extends from metathorax (T3) to caudal abdominal segment (AQ). 
True legs, prolegs and spiracles golden yellow. 

Fourth instar. Head: Diameter 3.8—4.0 mm. Color golden yellow. Body: Ground color 


168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


yellowish green. Length 26-29 mm, width 6 mm. All scoli golden yellow, dorsal and 
dorsolateral scoli with black tips on some spines. Light bluish gray mid-dorsal line extends 
from T3 to A8. Yellow subspiracular undulating fold extends from Al to A8. Small red 
patch with golden pinaculum in intersegmental area of sublateral surface just anterior 
of each proleg. Spiracles gold. True and prolegs yellowish green. 

Fifth instar. Head: Diameter 4.9 to 5.2 mm. Color green, adfrontal suture traced in 
brown. Body: Ground color green. Length 34 mm, width 9 mm. All scoli green, % or 
fewer of spines tipped with black. Thin red and black partially broken subspiracular line 
dorsal to broad white lateral line which may be edged in red on lower edge. Broad white 
and thin red lateral lines interrupted by lateral scoli. All three lateral lines (black, white 
and red) extend from posterior edge of T3 to A8. Small red patch with green pinaculum 
occur in intersegmental area of sublateral surface just anterior to each proleg and on A7. 
Dorsal surface green to bluish green, lateral and ventral surfaces green. Proleg shields 
reddish. Spiracles light brown. True legs green. 

Sixth instar. Head: Diameter 6.4-6.8 mm. Color and comments same as in previous 
instar. Body: Length 61 mm, width 12 mm. Ground color green. There is no noticeable 
difference between the fifth and sixth instar larvae other than overall size and a slight 
difference in the lateral lines. The black subspiracular line of the previous instar becomes 
red in the intersegmental areas and black only on the segment. A thin red line borders 
the ventral edge of the yellow lateral line. 


Key to the Last Instar Automeris Larvae of the United States 


1. Prolegs green; shields red or green. Prominent, continuous 
straight lateral yellow, white or red spiracular or subspiracu- 
lar line extends from metathorax or abdominal segment 1 
(Al) to A8 or beyond. Ventral surface primarily green ......... 2 
— Prolegs black, shields red or brown. White spiracular or sub- 
spiracular line broken or absent. Ventral surface black or red 


2. Spiracular or subspiracular lines white and red. Dorsal and dor- 
solateral abdominal scoli rosette type, shaft less than 5 mm 
Jonig' 365 so) Ua i ot nS A 3 

— Subspiracular line yellow, bordered dorsally by thin black line. 
Spiracular line absent. Dorsal and dorsolateral abdominal sco- 

li shafts slant anteriorly, shaft length 9-11 mm. (Arizona & 


News Mexico)". Cekiahah Naat hs Sie ey a A. randa Druce 
3. True legs pink or red. Prolegs green; shields pink or red. Spi- 
racular line.red 2.2.90 il 4 


— True legs green. Prolegs and shields green. Spiracular line ab- 
sent. Subspiracular line white, bordered by thin red or red 
and black line. (Arizona) A. iris hesselorum Ferguson 

4. Abdominal spiracles contained within red spiracular line. (Lou- 
isiana Ge WiISSIssijopol) eens A. louisiana Ferguson & Brou 

- Abdominal spiracles protrude from upper edge of red spirac- 
ular line. (various subspecies, widespread) A. io (F.) 

5. Dorsal abdominal area yellow with numerous black and light 
yellow lines extending length of larva. Two lateral white lines 
are interrupted by vertical yellow lines connecting lateral and 


VOLUME 39, NUMBER 3 169 


dorsolateral scoli. Scoli yellow and black. (New Mexico & 

MUESLCIMM UL GRAS) mtu es BE Acid We Peto Soo iel ta nd A. zephyria Grote 
— Dorsal abdominal area with numerous black, blue-gray or white 

lines extending length of larva. Diagonal white lines on lateral 

surface extend from base of dorsolateral scoli to lateral scoli 

on succeeding segment. Scoli blue-gray and black. (Arizona 

& New Mexico) A. cecrops pamina (Neumogen) 


Discussion 


To a greater extent than in any other state, the saturniid fauna of 
Arizona is significantly influenced by that of Mexico. Southern Arizona 
represents the most northern range of many Mexican plants, reptiles, 
birds, and insects (Lowe, 1964). The typically Mexican saturniid species 
such as Citheronia splendens sinaloensis Hoffmann, Eacles oslari 
Rothschild, Sphingicampa montana (Packard), Sphingicampa albolin- 
eata (Grote & Robinson), Adeloneivaia isara, Rothschildia cinctus 
Tepper, Automeris randa, and Automeris iris hesselorum usually oc- 
cur from 20 to no further than 80 km north of Sonora, Mexico. Other 
quasi-Mexican species [Anisota oslari Rothschild, Sphingicampa hub- 
bardi (Dyar), Hemileuca tricolor (Packard), Automeris cecrops pa- 
mina (Neumoegen), Eupackardi calleta (Westwood), and Agapema 
galbina anona (Ottolengui)] tend to be relatively abundant and with 
the exception of H. tricolor, occur from Arizona to Texas. 

The Mexican species that occur in Arizona can be placed into three 
categories: dependable resident, undependable resident, and tempo- 
rary resident. Citheronia splendens and Eacles oslari are examples of 
dependable residents. They are not necessarily common but represent 
species that are found each year. Undependable residents include 
Rothschildia cinctus, Sphingicampa albolineata, and Sphingicampa 
montana. These are permaneut residents which are infrequently cap- 
tured, because they occur in isolated areas, generally have low popu- 
lation levels, and lack biological and habitat data. For example, seven 
blacklight stations were set up in the southern branch of Pena Blanca 
Canyon and operated for two consecutive nights. Each station had two 
to three 15 watt blacklights and was separated from the next station 
by 75 to 100 m. One R. cinctus and two S. montana were captured; 
both montana were taken at the same light. On the third and fourth 
nights a mercury vapor light was operated, and nearly a dozen mon- 
tana were captured. If each station had been operated by a different 
collector, most might have been assumed that neither S. montana nor 
R. cinctus were present. Collecting techniques, location, experience, 
persistence, and chance interact to influence the collector’s perception 
of species abundance and distribution. 


170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Automeris randa is probably best described as an unpredictable 
resident. The fact that randa occurs over a wide area within the Pe- 
loncillo Mountains of Arizona and New Mexico suggests it is well es- 
tablished. This species probably remained uncollected due to the iso- 
lation and inaccessibility of its habitat. During a trip to Guadalupe 
Canyon we traveled for over 20 miles on unmarked dirt roads and 
after arriving, found that only vehicles with high road clearance could 
actually enter the canyon. 

Automeris iris hesselorum and Adeloneivaia isara are the most re- 
cent, but perhaps not the only, examples of temporary Arizona resi- 
dents. Specimens of Citherona mexicana (Grote & Robinson) and 
Sphingicampa albolineata were labeled “Cochise Co., Arizona” and 
Hylesia coinopus Dyar as “So. Arizona/Poling” and are treated as 
mislabeled or with suspicion. The ephemeral nature of these species 
may be influenced by seasonal variation in the weather of northern 
Sonora and southern Arizona. Perhaps a succession of favorable winters 
and summers allows the temporary /ephemeral species to become es- 
tablished in Arizona, but periodically, climatic conditions exceed their 
tolerances, causing local extinction. Based on the observation that S. 
albolineata is now known to be a resident species in Cochise Co., the 
ephemeral nature of A. iris hesselorum and A. isara, and the recent 
discovery of A. randa, the past occurrence of Citheronia mexicana 
and Hylesia seem more likely. 


ACKNOWLEDGMENTS 


I wish to thank Peter Jump, Steve Prchal, and Ron Wielgus for their observations, and 
Steve McEflresh, Scott Meredith, Tom Carr, and Jim Tuttle for accompanying the author 
on Arizona trips, and for sharing ova and observations. I would also like to thank Ann 
McGowan-Tuskes and Mike Collins for their comments on the manuscript. 


LITERATURE CITED 


FERGUSON, D. C., in Dominick, R. B. et al. 1972. The moths of America north of 
Mexico, Fasc. 20.2B, Bombycoidea (in part). 
Lowe, C. H. 1964. Arizona’s natural environment. Univ. Ariz. Press. 136 pp. 


Journal of the Lepidopterists’ Society 
89(3), 1985, 171-176 


COURTSHIP AND OVIPOSITION PATTERNS OF TWO 
AGATHYMUS (MEGATHYMIDAE) 


DON B. STALLINGS AND VIOLA N. T. STALLINGS 
P.O. Box 106, 616 W. Central, Caldwell, Kansas 67022 


AND 


J. R. TURNER AND BEULAH R. TURNER 
2 South Boyd, Caldwell, Kansas 67022 


ABSTRACT. Males of Agathymus estelleae take courtship sentry positions near ten- 
eral virgin females long before the females are ready to mate. Males of Agathymus 
mariaeé are territorial and pursue virgin females that approach their territories. Ovipo- 
sition patterns of the two species are very similar. Females alight on or near the plants 
to oviposit and do not drop ova in flight. 


Few detailed observations of the courtship and oviposition of the 
skipper butterflies in natural environments have been published. For 
the family Megathymidae Freeman (1951), Roever (1965) (and see 
Toliver, 1968) described mating and oviposition of some Southwestern 
U.S. Agathymus, and over a hundred years ago (1876) Riley published 
an excellent paper on the life history of Megathymus yuccae (Bois- 
duval & LeConte) which included data on oviposition of the female; 
otherwise, only the scantiest comments have been made. C. L. Rem- 
ington (pers. comm.) and others tell us that there is a significant pos- 
sibility that the Hesperioidea are less closely related to the true but- 
terflies (Papilionoidea) than to certain other Lepidoptera and even that 
the Megathymidae may not be phylogenetically linked to the Hesper- 
iidae. For several years we have been making on-the-scene studies of 
these two aspects of megathymid behavior, both for their interest in 
understanding the whole ecology of these insects and for their possible 
reflection on higher relationships. In this first paper we are presenting 
our findings for two close relatives in the genus Agathymus. 

In 1976 the four of us took advantage of an opportunity to watch a 
number of courtship sequences of Agathymus estelleae (Stallings & 
Turner) and their pattern of ovipositing. Most of these observations 
were made on 7, 8, 12, and 14 September, 16.5 km north of Saltillo, 
Coahuila, Mexico on Highway 57 at an elevation of approximately 
1380 m. The females emerged from 0800 to 0930 h (CDT) and in the 
wild, crawled up on a leaf of their food plant as their wings expanded 
and hardened. Shortly after the females commenced to emerge, males 
appeared and flew by the female, often as close as 80 cm. The first 
male to locate a female would then perch on a leaf of the food plant, 
Agave lecheguilla Torr., or on a stone or small shrub about 8 m or less 


we JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


downwind from the female; other males would fly by the female and 
would be pursued by the first male, who would chase them out of his 
“territory.”” The sentry site of the first male appeared to be rather 
small, as subsequent males would take up positions downwind from 
the female as close as 5 m from the first male. There tended to be 
three males in attendance by the time of the maiden flight. Butterflies 
would fly through the area and would not be pursued by the males 
except for dark skippers with white fringes, which the males evidently 
mistook for other estelleae. After the female’s wings had expanded 
fully, she would rest on the leaf with her wings folded so that the apex 
of the forewings touched. The males, on the other hand, rested with 
their forewings apart, and their hindwings dropped down almost per- 
pendicular with their body. We called this the male launching position, 
as they were able to take off in full flight immediately. 

Two to three hours after a female emerged she would make her 
maiden flight, which seemed a long period to us. She would rise from 
her resting position in a circular flight and fly downwind very rapidly 
in an undulating manner some 2 to 5 m above the ground. In nearly 
every instance she flew directly over the male who had been first to 
establish a sentry position. The first male would rise above and just 
behind to meet her as she flew over his position dipping down towards 
her so that he appeared to touch her. She would quickly drop down 
to or near a food plant, followed closely by the male. All of this hap- 
pened within a few seconds, in which the female had not traveled over 
40 m from her original resting site. If there were other males watching 
the female, they joined in the pursuit of her. Usually as the female 
and first male came to rest, they were in copulation within 8 or 4 sec. 
If there were other males they would alight by the female and try to 
mate with her. If none of the males had succeeded in mating with her 
in about 7 sec, she would fly off in a straight line pursued by all of the 
males. We were never successful in observing what happened when 
she came to rest a second time. On three occasions, if the first male 
was successful in mating, the other males flew away but returned with- 
in 5 to 10 sec and alighted within 15 to 20 cm of the copulating pair. 
Within a few seconds the unsuccessful males again flew away and then 
returned within 5 to 10 sec but would alight 60 to 100 cm from the 
copulating pair. Again, the unsuccessful males would fly away and 
return shortly, this time alighting about 2 m away from the pair. After 
a final brief inspection, the unsuccessful males would fly away and not 
return. The mating pair would remain in copulation from 66 min to 
five hours. We had no difficulty in moving them into a wire cage, so 
that we could recover the ova to be laid later. On two occasions we 
happened to flush virgin females before they were ready to mate. Their 


VOLUME 39, NUMBER 3 173 


flight was in a straight line to another plant or bush, and although there 
were males around, none of them pursued these females. 

On 7 September a pair was found in copulo at about 1430 h, clinging 
to the underside of a lecheguilla leaf; when they flew away, the female 
appeared to be flying and the male dangling. This copulation had 
presumably started much earlier, because four observed couplings took 
place at 1050, 1118, 1120, and 1155 h. 

While vision is undoubtedly a major part of the courtship process, 
we assume that the female emits a pheromone shortly before or as she 
makes her pre-mating flight. On one morning the wind shifted after a 
male had established his sentry position. We noted with interest that 
he maintained his position even though he was then upwind from the 
female. When the female took off on her mating flight she flew down- 
wind, with the result that the first male was an unsuccessful suitor, 
because a downwind male reached her before he did. 

One observation day was very hot, and from noon until 1600 h we 
saw no flight activity among the estelleae, although there were other 
butterflies flying in the area. Shortly after 1600 h some scattered clouds 
appeared along with some female estelleae. Each of the females pro- 
ceeded to oviposit by alighting on a leaf of the food plant with wings 
completely closed and then dropped an ovum that fell to the base of 
the plant where it might lodge among the leaves or bounce out on the 
ground. This took from four to seven seconds. We were wondering 
why the first females did not drop more than one ovum at a time, 
when we were accommodated by a female which proceeded to drop 
five to seven ova at one location, without flying. A few days later we 
again watched females lay as many as five ova in one sitting. We noted 
that after a female dropped five to 10 single or multiple ova she would 
alight on a rock or bush and rest before proceeding with her ovipos- 
iting. Since in many years of field work we had seldom found larvae 
in juvenile plants or plants located in the shade of a bush or rock, we 
had always supposed that the females were very selective as to where 
they placed their ova. However, we were in error, as the females were 
indiscriminate as to the size or location of the plants where they dropped 
their ova. We suspect that there are more predators in the juvenile 
plants or plants under bushes and that the ova or the small larvae had 
a lesser chance to survive there. 

At about 1630 h the cloud cover usually became heavier, and the 
temperature dropped slightly. Various butterflies in the vicinity sought 
shelter, but the estelleae females continued their activities. Once a 
slight breeze came up, and we could smell moisture. Evidently, the 
estelleae recognized the oncoming shower, for they immediately sought 
shelter in plants and bushes and on the downwind side of small rocks. 


174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A minute or two later it began to rain, and it was evident that several 
of the females had not picked a good site for shelter as they com- 
menced flying about seeking a more sheltered area. 

One of the females that we kept for ova count produced 95 ova the 
first day, 23 the second day, 10 the third day, 23 the fourth day, 4 the 
fifth day, 14 on the sixth day, and 8 on the seventh day; when she died 
she had 8 ova left in her body for a total of 180 ova. Ova of this species 
vary in color from green through a sand color; all of those laid on the 
last two days by this female were green. 

Two of the authors (DBS and VNTS) conducted a series of obser- 
vations 5 and 6 October 1983 in the Guadalupe Mountains near Carls- 
bad, New Mexico for the purposes of determining the courtship and 
Ovipositing patterns of Agathymus mariae (Barnes & Benjamin), a 
species rather closely related to A. estelleae. 

Females emerged from their pupae from 0900 to 1030 h (MST). The 
males emerged some 30 min earlier than the females. A newly emerged 
female would crawl up a leaf of the plant on its inner side, approxi- 
mately one-third of the distance from the tip of the leaf. It took about 
2 to 3 hours for the wings to expand and harden. 

Males that had emerged the prior day or earlier patrol the area 
around the food plants (Agave lecheguilla). Males that emerged earlier 
in the day eventually join in the patrol. The patrolling on the part of 
the males consists of flying back and forth over the food plants. It is 
interrupted by the males alighting on small rocks or directly on the 
ground in an open area where they can see and be seen; they would 
remain there from 1 to 10 min and then resume their patrol. The 
patrol flight was usually within 1 m of the ground. As they repeated 
their patrol and alighting procedure, they would sometimes alight on 
the ground, where they had been, and at other times would alight in 
a new area. The selection by a male of a new area within which to 
alight may be dictated by the failure of a female to fly over the pre- 
viously selected site. In defending territory a male would pursue another 
male who entered his territory. They would often fly 6 to 10 m up into 
the air. 

Males would often fly by a teneral female within 30 to 40 cm without 
any indication that they recognized her presence. 

It appears that mariae males alight in an open area and depend on 
a female finding them by flying over them. This is in contrast to es- 
telleae where the male establishes a sentry position in relation to a 
specific female which he will pursue when she makes her first flight. 
Thus, while estelleae and mariae have different courtship patterns, the 
males of each establish a position during the courtship period of each 
day and defend it against other males. 


VOLUME 39, NUMBER 3 175 


We noted a number of day-flying saturniid moths (Pseudohazis) 
flying about, which we sometimes mistook for an Agathymus, but the 
male mariae evidently had no such difficulty. 

Around noon a female would take off on her first flight. She would 
circle in a fluttering manner around the area where she had been 
resting. Her flight was usually not over 70 cm above the ground, and 
the radius of the circle was about 8 or 9 m. This circle nearly always 
covered an area that would have at least one male temporarily on the 
ground. When the male sighted the female, he would take off in pur- 
suit of her. As the male approached the female, she would drop to the 
ground, alighting on a rock or a leaf of the food plant where he would 
join her; within 5 sec they would be in copulation. The duration of 
copulation varied from 48 min to several hours. 

After copulation the fertilized female would remain at rest until 
1600 or 1700 h, at which time she would commence ovipositing. We 
think the period of day when females oviposit is determined by tem- 
perature, because captive females in protected areas where the tem- 
perature was less than in more exposed areas would commence ovi- 
positing shortly after mating. A female would fly in a fluttering manner 
to a food plant where she would alight on one of the outer leaves and 
drop an ovum. Some females dropped a single ovum while others 
dropped two ova. We suspect that some females drop more than two 
ova at a single stop, although we did not observe any doing this. 

Since the females that we observed alighted on the outer leaves of 
the food plant, all of the ova fell on the ground. We were able to 
recover ova that we saw laid. The female would usually take a short 
rest after depositing 8 to 5 ova. 

The three couplings that we observed occurred at noontime, specif- 
ically at 1155, 1210 and 1242 h. 

Our observations of Agathymus carlsbadensis (Stallings & Turner) 
in this same general area of the Guadalupe Mountains was inconclu- 
sive. We think the courtship pattern is very similar to that of A. mariae. 
The ovipositing pattern appears to be entirely different. We never did 
observe a female alighting on a plant to oviposit. At least five different 
times we observed females hovering over or near their food plant. We 
suspect that they were ovipositing from this hovering position, but we 
were unable to recover any ova. 

In the 1940’s we corresponded with W. P. Medlar, a collector in 
California who advised us that he had observed Agathymus stephensi 
(Skinner) ovipositing. According to him the female hovered over or 
near the food plant and flipped the ova towards the plant. Freeman 
reported the same thing in 1951 for Agathymus aryxna (Dyar) and 
Agathymus evansi (Freeman). In 1951 when Freeman’s paper was 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


published the name neumoegeni was being mistakenly applied to 
aryxna. He followed this mistake. Correspondence that we received 
from Freeman at the time of his observations mention only evansi as 
ovipositing from a hovering position; no mention was made of aryxna. 
Roever (1965) reported that he had not observed females flipping their 
ova into the plant while hovering over or near a plant; however, he 
noted that both of the species alight on or near the plant when ovi- 
positing. He reported that he had observed ovipositing by the female 
while on or near a food plant. He made the same observations for 
Agathymus neumoegeni (Edwards), A. polingi (Skinner), A. baueri 
(Stallings & Turner), A. freemani (Stallings & Turner). It is evident 
that more observations are needed in order to reconcile these divergent . 
reports. 


ACKNOWLEDGMENT 


We wish to thank Dr. Charles L. Remington for his advice and assistance in our studies 
of the Megathymidae and particularly his advise in the drafting of this paper. 


LITERATURE CITED 


FREEMAN, H. A. 1951. Notes on the Agave feeders of the genus Megathymus. Field 
& Laboratory 19:26-32. 

RILEY, CHARLES V. 1876. The yucca borer.—Megathymus yuccae (Walker). Eighth 
Ann. Rept. Noxious, Beneficial and Other Insects of the Station of Missouri, pp. 169- 
182. 

ROEVER, KILIAN. 1965. Bionomics of Agathymus. J. Res. Lepid. 3:103-120. 

TOLLIVER, MICHAEL. 1968. Apparent partial courtship between Megathymus yuccae 
coloradensis and M. streckeri (Megathymidae). J. Lepid. Soc. 22:177-178. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 177-186 


BIOLOGY OF THE HALF-WING GEOMETER, 
PHIGALIA TITEA CRAMER (GEOMETRIDAE), AS A 
MEMBER OF A LOOPER COMPLEX IN WEST VIRGINIA! 


LINDA BUTLER 


Division of Plant and Soil Sciences, P.O. Box 6108, 
West Virginia University, Morgantown, West Virginia 26506-6108 


ABSTRACT. Field and laboratory studies were conducted during 1983 in two coun- 
ties of eastern West Virginia where forest defoliation by a looper complex (Geometridae) 
had been heavy the previous two years. Phigalia titea Cramer, dominated, making up 
77-94% of feeding larvae; Erannis tiliaria (Harris), P. strigateria (Minot) and Alsophila 
pometaria (Harris) were also present. Adult P. titea were found in the field from 17 
March to 26 April, eggs from 17 March to 3 May and larvae from 3 May to 15 June. 
Descriptions of oviposition sites and eggs are given; females were found to contain a 
maximum of 1447 eggs. At constant 24°C P. titea larvae required a mean total of 28 
days to mature through five instars when fed on sugar maple leaves, but larval growth 
rates were found to vary with host plant species. Descriptions of the five larval instars 
are given. 


As defoliators of hardwood forests numerous species of native geo- 
metrids produce either consistent but little noticed damage or sporadic 
but significant damage during outbreaks in eastern hardwood forests. 
Outbreaks may consist primarily of a single species or represent a 
complex of geometrid defoliators. 

During the spring of 1981 and 1982, approximately one million acres 
of hardwoods in eastern West Virginia were defoliated each year to 
varying degrees ranging from 20-100%. The West Virginia Depart- 
ment of Agriculture assessed the looper infestation as causing more 
damage to hardwoods in one year than all other defoliators in the state 
have in the past 20 years. Loopers have been the most obviously de- 
structive forest insects in West Virginia in recent times, with numerous 
reports of tree mortality following heavy defoliation (Anon., 1983). 
While various species were suggested as comprising the defoliator com- 
plex, no detailed observations had been made (Anon., 1981, 1982). 

The extensive defoliated acreage and resulting mortality of hard- 
woods, especially oaks and hickories, justified a more detailed study on 
the looper complex in West Virginia. Sampling of larval populations 
showed four looper species to be present at all study sites: Phigalia 
titea Cramer, P. strigateria (Minot), Erannis tiliaria (Harris) and Al- 
sophila pometaria (Harris). In all samples P. titea made up the ma- 
jority of the looper population, ranging from 77% to 94%, depending 
on the site. While some information on P. titea is available (Baker, 


! Published with the approval of the West Virginia University Agricultural and Forestry Experiment Station as 
Scientific Article #1884. 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1972; Talerico, 1968), it is often of a superficial nature or contains 
information contrary to observations made during this study. The out- 
standing exception is the fine study of adults by Rindge (1975). 


METHODS AND MATERIALS 
Study Area 


Three sampling sites were selected in West Virginia. Two were in 
Cacapon State Park in Morgan County of West Virginia’s Eastern Pan- 
handle within the most heavily defoliated region; the third site was 
near Elkhorn Mountain at the border of Grant and Hardy counties on 
the southern edge of the affected region. The vegetation at all sites is 
often referred to as oak-hickory-pine, although originally chestnut was 
a dominant species. Both study areas are dry upland sites consisting in 
part of steep shale slopes. 

The two Cacapon State Park sites were on Cacapon Mountain. The 
site designated Batt Picnic Area (Batt PA) was east facing at an ele- 
vation of 381 m and had complete looper defoliation in 1982. The 
second site, designated Cacapon Overlook (Cac. OL) was along both 
sides of a north/south ridge at an elevation of 701 m and suffered 
about 25% looper defoliation in 1982. The Elkhorn (Elk.) study area 
was on Getz Mountain, 1-2 km south of Elkhorn Mountain, at an 
elevation of about 732 m. During 1982 the Elkhorn area also received 
about 25% looper damage. 


Field Collection and Description of Looper Life Stages 


Sampling for all looper species was initiated with adult observations 
on 17 March 1983 at Cacapon State Park and 24 March at Elkhorn 
Mountain. Samples were taken at 6- to 8-day intervals through 15 June, 
at which time most larvae had moved into the soil for pupation. 

During the weeks of adult Phigalia activity, collections were made 
and relative numbers and locations of males and females were noted 
during timed walks through each study site. Males were determined 
to species through genitalia dissection; females were collected for de- 
termination of species and fecundity. Fecundity was studied by two 
methods: (1) dissection of field collected females and (2) caging of 
females on dead twigs for oviposition, counting deposited eggs and 
then dissecting the post-ovipositional females to count residual eggs. 

Phigalia eggs were observed in the field and developmental color 
changes noted on a weekly basis. Eggs on dead twigs or other vege- 
tation were collected from 17 March through 26 April, were held 1 to 
4 weeks at 4°C, then placed at room temperature to hatch; viability 
was determined. 


VOLUME 39, NUMBER 3 179 


TABLE 1. Potential egg counts from dissected female Phigalia titea collected at var- 
ious dates and study sites in 1983. Sample size given in parentheses. 


Site Date No. eggs/female Range 

Batt PA III.17 (40) 309 21-884 
III.24 (5) 215 11-454 
EV ein 7) 48 9-114 
VAS (C7) 23 8-51 
VE2OF(S) 29 7-54 

Cacapon OL IJI.24 (1) 1364 
III.31 (6) 542 395-836 
IV.14 (6) 859 288-1409 

Elkhorn I]J.24 (2) 910 643-1177 
III.31 (6) 775 177-1447 


At each sampling date between larval hatch and pupation, foliage 
samples were collected with pole pruners and placed in plastic bags 
for transport to the laboratory. Samples of 100 to 200 larvae were taken 
from the pruned vegetation and preserved for determination of species, 
and in the case of P. titea, to determine instar composition and to 
prepare larval descriptions. 


Larval Development 


To determine intervals between larval molts, newly hatched larvae 
from the egg viability study were placed in groups of 10 on leaf clusters 
of host plants in petri dishes and held at 24°C. In one experiment, 200 
first instar larvae were reared on sugar maple; one group of 100 set up 
27 April, a group of 20 set up 6 May and a group of 80 set up 11 May. 
In a second experiment begun 4 May, growth rates of larvae were 
compared when fed on red oak and red maple; 50 larvae were reared 
on each food plant. 


RESULTS AND DISCUSSION 
Fecundity 


Most of the P. titea females collected from Batt PA for dissection 
were already ovipositing, and the resultant potential egg count may 
have been lower than the number of eggs which they were capable of 
producing. The expected trend of lower potential egg numbers per 
female with increasing time into the season is shown in Table 1. Fe- 
males collected at Batt PA from 7 to 20 April were depleted. 

Egg counts of dissected females from Cac. OL and Elk. were con- 
siderably higher than oviposition rates previously reported for P. titea; 
Talerico (1968) reported that several caged females produced from 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Dissected ovaries of a Phigalia titea containing 1364 eggs. 


127 to 189 eggs each. Data show that dissection of preovipositional 
females gives an accurate indication of reproductive potential since 
90% of eggs were laid in the laboratory cages (Table 2). The highest 
egg count from a dissected female was 1447. Figure 1 shows dissected 
ovaries of a P. titea female with 1364 eggs. The four ovarioles per 
ovary contained the following egg numbers: 166, 164, 193, 157, 184, 
185, 153, 162. 


Oviposition Sites and Egg Descriptions 


Eggs were generally laid on dead twigs, singly or in clusters, with 
the cluster configuration and size depending upon structural features 
of the oviposition site. Common oviposition locations were under loose, 
flaking bark of dead twigs of maples (Acer spp.), oaks (Quercus spp.), 
black gum (Nyssa sylvatica Marsh.), common witch hazel (Hamamelis 
virginiana L.), flowering dogwood (Cornus florida L.), grape (Vitis 
spp.) and around bark cracks of hickory (Carya spp.) twigs. Many 
clusters were under loose bark at crotches of dead twigs. Other loca- 
tions included deep splits in dead twigs, bostrichid burrows, frass of 
shallow buprestid or cerambycid burrows, exposed face of girdled twigs, 
empty chorions of Saturniidae, small Lepidoptera pupae or cocoons, 
spider egg sacs, and dead crumpled leaves of composites from the 
previous year. One of the most interesting locations was inside empty 
chorions of previous year’s eastern tent caterpillar, Malacosoma ameri- 


VOLUME 39, NUMBER 3 181 


TABLE 2. Egg counts for female Phigalia titea allowed to oviposit in the laboratory 
and subsequently dissected. Sample size given in parentheses. 


# eggs/dissected 
fenaate Mean 
a a a 2 pe Se Se total / 
Site Date Mean Range Mean Range female Range 


Cacapon OL IV.7 724 496-1035 65 3-243 789 551-1266 


# eggs laid 


(8) 
IV.14(3) 686 242-1022 26 811-49 712 260-1133 
IV.20(5) 664 280-1128 40 3-123 704 299-1251 
Elkhorn IV.7 (5) 394 72-545 36 7-83 4380 155-597 
IV.14(9) 601 41-1140 49 2-278 650 60-1198 
(5) 


337 110-432 53 09-147 390 115-516 


canum (Fab.), egg clusters (J. E. Weaver, pers. observ.). Diameters of 
65 randomly collected egg-bearing twigs of various host species aver- 
aged 7.2 mm (range 2-15 mm) at the oviposition sites. 

Eggs were slightly roughened by reticulate sculpturing, as previously 
described by Forbes (1948). The shape was oval with one end of the 
egg more flattened or broadly rounded and the opposite end more 
conical; heaviest sculpturing was at the broad end. Measurement of 80 
eggs collected at both sites at Cacapon State Park on various dates 
averaged 0.905 (0.858-0.990) mm long and 0.521 (0.495-0.561) mm 
wide: These sizes were only about half that reported for P. titea eggs 
in Virginia (Talerico, 1968). 

When first deposited, eggs were greenish yellow to yellow in contrast 
to an earlier description by Talerico (1968). Further color changes 
reflected embryonic development, the rate of which is temperature 
dependent. Within 2 to 8 days at 24°C, eggs began developing a salmon 
pink color which first appeared at the blunt end of the egg. The eggs 
gradually became darker pink over a period of several days, and when 
approaching maturation, dark red to black spots appeared at the blunt 
end. At about 24 h prior to hatch, the eggs appeared dark purple as a 
result of the dark head capsule and body of the larva being visible 
through the lavender chorion. At this stage, microscopically, the larva 
was seen curled inside with head and posterior end meeting at the 
blunt end of the egg. The pale spiracular stripe of the larva was easily 
seen. At constant 24°C, development time from oviposition to hatch 
required 7 to 8 days. 

Larvae eclosed by chewing an irregular hole through the blunt end 
of the egg. Empty chorions were an iridescent lavender and were easily 
observed in the field. Eggs parasitized by Telenomus alsophilae Vi- 
ereck (Hymenoptera: Scelionidae) appeared chocolate brown at the 
time adjacent unparasitized eggs were hatching. In the laboratory, 
parasite adults did not eclose until about 13 to 15 days after P. titea 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Comparison of developmental time for Phigalia titea larvae reared on 
sugar maple,’ red oak’ and red maple. 


Time in instar (days) 


Mean Range 
Instar Sugar maple Red oak Red maple Sugar maple Red oak Red maple 
1 4.30 4.24 4.50 3-5 3-5 3-5 
2 3.02 3.18 3.18 2-5 2-4 2-4 
3 3.50 2.74 3.70 2-5 2-4 2-5 
4 3.43 3.36 4.30 2-6 2-4 3-6 
a) 7.66 6.92 7.24 6-11 6-8 6-9 
Prepupa 6.90 5.88 7.06 4-10 5-7 6-9 
Total 28.41 26.32 29.98 21-40 20-31 24-36 


2 Based on 200 larvae. 
> Based on 50 larvae. 


larval hatch. Chorions of parasitized eggs retained a brown color, thus 
allowing easy evaluation of percent parasitism in a field situation. 

Percent hatch of 12,855 eggs randomly collected at the three study 
sites averaged 94.4. Some eggs which failed to hatch had obviously 
suffered mechanical injury at collection; many eggs remained dark 
yellow as if no embryonic development had occurred, while others 
were dark pink. Dead larvae were observed in some eggs. Parasitism 
was low in all egg collections and, at most sites, did not appear to 
contribute significantly to mortality. 


Larval Growth Rates 


As reported by Talerico (1968) and confirmed in this study, P. titea 
has five larval instars. Growth data for 200 larvae reared from eggs 
which hatched on three different dates are combined and given in 
Table 8. It was obvious from this evaluation that time spent per instar 
was related to developmental stage of host plant foliage. Larvae which 
hatched on 27 April were reared on the flowers and very young leaves 
of sugar maple, larvae from 6 May on young to moderately aged leaves 
and larvae from 11 May primarily on mature leaves. Greater length 
of time was required between lst instar and prepupation with increas- 
ingly older foliage, e.g. larvae set up on 27 April matured on the 
average 6 days faster than larvae set up on 11 May. This difference is 
not apparent in Table 3, however, since all data are combined. 

As expected, larvae reared on different host plants develop at dif- 
ferent rates. The period from hatch to pupation was about 26 days for 
red oak and about 30 days for red maple. Field observations during 
the 1983 season substantiated this finding. At any point during the 
larval season, later instar larvae were always on oak, hickory and birch 
and earlier instars on dogwood, maple, blackgum, witch hazel, and 
most other hosts. 


VOLUME 39, NUMBER 3 183 


Larval Description 


Talerico (1968) made only brief reference to larval color patterns. 
Descriptions of color patterns of each instar from the populations with- 
in the West Virginia study areas are given below. Head capsule widths 
are the average of 50 specimens per instar. 


Instar 1. Head capsule (0.8329 mm) medium to dark reddish brown with paler frons. 
Ground color of body of newly hatched larvae slate grayish black. Dorsal pinnaculae 
dark, dull yellow; pair of small pale yellow middorsal pyramidal streaks at posterior 
margin of each segment from mesothorax through abdominal segment 6. Spiracular stripe 
from creamy white to pale yellow, extending almost continuously from prothorax through 
abdominal segment 6. Subspiracular area brownish to yellowish brown. Caudal shield 
and anal prolegs medium brown to dull yellow-brown. Late Ist instar larvae appear 
much more pale, with medium olive ground color, after intersegmental areas are exposed 
with larval growth. Particularly prominent just prior to molt is dull yellow-brown cervical 
area which is protuberant and dwarfs head capsule. In mature Ist instars, caudal, cervical, 
and proleg sclerites medium brown and very evident against paler body. 

Instar 2. Head capsule (0.621 mm) dark reddish brown to black with pale maculations. 
Basic body ground color greenish brown to slate black. Paired cervical shields small but 
prominent. Pair of fine, indistinct white dosal stripes down length of body but discontin- 
uous between segments. Pinnaculae dark. Spiracular stripe diffuse and faint, but present; 
fades out on abdominal segment 7. Secondary setae present but sparse on body, located 
primarily above spiracular line. 

Instar 3. This is the first instar that begins to develop a striping pattern approaching 
that of mature larvae. These larvae, however, are very dark; mature individuals appear 
shiny black, with striping being evident only under magnification. 

Head capsule (1.05 mm) black with prominent grayish white mottled areas. Fine 
irregular grayish white striations down length of body. Paired dorsal stripes filled with 
small orange spots on posterior margin of each segment. Pinnaculae black. Diffuse orange 
coloration along spiracular area most prominent on protuberances of abdominal segments 
1 and 5; lateral orange areas of thoracic segments and abdominal segment 6 reduced. 
Caudal and cervical areas black with white mottling; venter black. Secondary setae more 
numerous than in previous instar including some below spiracular line. 

Instar 4. Head capsule (1.87 mm) black and white mottled with a higher proportion 
of white than in previous instar. Frons mottled, clypeus white. Two pairs of black irreg- 
ular dorsal stripes, orange-filled. Supraspiracular stripe a pair of black irregular lines, 
white-filled and flanked with orange. Spiracular area with pair of black irregular lines, 
white-filled, running just above and below spiracle. Area in immediate vicinity of spiracle 
on each segment prominently suffused with orange. Setae SD1, L1 and L2 within this 
orange area and each on separate black chalaza. Spiracular chalazae of second abdominal 
segment most prominent; abdominal segment 8 with pair of prominent chalazae forming 
dorsal hump. Subspiracular stripe of double irregular black lines, grayish-white-filled. 
Shields mottled black and white. Legs black; abdominal sternites 7 to 8 white. 

Instar 5. Head capsule (2.83 mm) white with distinct black maculations. Body ground 
color pale lavender-gray with pairs of fine irregular black lines, orange-filled dorsally, 
pale gray-filled subdorsally. Orange patterns, chalazae, shields and venter as in instar 4. 
Peritremes black; spiracular valve off-white. Basic appearance of this larva paler with 
striping more prominent than in previous instars. 


P. titea Pupae 
The pupa of P. titea is illustrated by Talerico (1968). During the 
current study, measurements were made of 80 pupae; mean length was 
13.45 mm and range was 11-15.5 mm. Length was not related to sex, 
but female pupae were characteristically stouter. 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Life History 


Adult P. titea emerged from the soil where they overwintered as 
pupae and climbed vertical surfaces. Males eclosed several days to a 
week before females and were most commonly seen resting on tree 
trunks. Females climbed tree trunks of a wide range of sizes and re- 
mained there for a period of hours to a day or two. Females (and thus 
eggs) were rarely found on small dead trees. Mating most commonly 
occurred on tree trunks, after which females climbed upward to locate 
suitable oviposition sites. 

Male P. titea were first observed flying at Berkeley Springs, WV, 
near Cacapon State Park on 8 March. At Batt PA on 17 March, males 
were numerous and resting on tree trunks. The total numbers of fe- 
males appeared relatively less than males, because of their virtual 
winglessness and related cryptic appearance and, because their behav- 
ior rendered them more difficult to observe; 40 females were found in 
a 2 h walk. Most of the Batt PA females were on dead twigs, but some 
were still emerging from the soil and beginning to climb tree trunks. 
Of the few eggs present on the twigs, about 99% were yellow. The 
number of adults seen on 24 March was similar to that of the previous 
week; numbers of observed females and males began to decline by 31 
March and continued to decline markedly each week. The last male 
was observed at Batt PA on 14 April and the last female on 26 April. 

A few males were seen at Cac. OL on 17 March and at Elk. on 24 
March. Females were observed for the first time on the latter date at 
both sites. Numbers of adults continued to increase at Cac. OL and 
Elk. through 7 April, then sightings began to decline on 14 April. No 
males and low numbers of females were seen on 26 April. Adult pop- 
ulations of P. titea were markedly lower at Cac. OL and Elk. than at 
Batt PA. At peak female activity, an average of 8 to 10 were observed 
in a 2 h walk. 

A few eggs were seen at Batt PA on 17 March with more than 99% 
of these being yellow. The first ovipositing females were observed at 
Elk. on 24 March and at Cac. OL on 7 April. By the later date, about 
10% of eggs at Batt PA were pink and females at Elk. and Cac. OL 
were reaching peak oviposition activity. On 14 April, about 90% of the 
eggs at Batt PA were pink, while at other sites only about 20% had 
developed pink coloration. By 26 April, 95% of the eggs at Cac. OL 
and Elk. were pink. 

Egg hatch began in early May and by 3 May was about 85% com- 
plete at Batt PA; remaining eggs were in the purple stage, indicating 
hatch would occur within about 24 h. At Batt PA, Ist instar larvae 
were very evident, as they hung on silk lines below the dead twigs on 


VOLUME 39, NUMBER 3 185 


TABLE 4. Instar composition (% of population) of Phigalia titea taken at study sites 
between 3 May and 8 June. 


Study site 
Date Instar Batt PA Cacapon OL Elkhorn 

V.3 1 87 90 100 

2 13 10 
V.10 if 6 73 94 
2 87 26 6 

3 Ji ji 
Veh7 1 0 40 0 
2 4 52 82 
3 09 7 eZ, 
4 37 1 1 
V.24 2 0) 13 3 
3 2 67 Tl 
4 28 if 26 
5) 70 1 0 
V.31 3 0 4 3 
4 a 78 52 
3) 98 18 45 
VL8 4 8 if 
fs) 92 93 


which eggs had been laid. Ballooning actively occurred, with the larvae 
riding air currents into trees which were just beginning to leaf. It 
appeared that larvae arrived at potential host trees randomly by means 
of wind activity, but they showed preference for hosts by either settling 
quickly and beginning to feed or continuing to balloon if the initial 
host was not suitable. Considerable larval mortality probably occurred 
at this time. 

On 8 May at Batt PA, some larvae were beginning to feed with 
damage being initially in the form of pinholes and skeletonizing. On 
this date at Cac. OL, egg hatch was about 20% complete but was just 
beginning at Elk. No feeding was evident at these sites. 

Despite the fact that eggs were laid over a period of several weeks 
at each site, egg hatch at any one site occurred relatively in synchrony 
due to effect of low temperatures on earlier laid eggs. Progression of 
P. titea larval development for each of the study sites is given in Table 
4, Differences in altitude and temperature at the study sites produced 
developmental events, resulting in adult emergence to pupation being 
one to two weeks earlier at Batt PA than at the other two sites. Dif- 
ferences in developmental time also occurs depending on orientation 
of slope. 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


I wish to thank Superintendent Philip Dawson and his staff of Cacapon State Park for 
their cooperation during this study. Particular thanks go to Mr. and Mrs. Fred Riggleman 
of Dorcas, WV for allowing me to conduct a portion of this research on their land at 
Elkhorn Mountain; their cooperation and interest were greatly appreciated. For assis- 
tance in laboratory studies I am indebted to Vicki Kondo, Terry Stasny and Beth Cahape. 
I also thank James W. Amrine, David E. Donley, John E. Hall and Joseph E. Weaver 
for comments on the manuscript. 


LITERATURE CITED 


ANONYMOUS. 1981. Forest insect and disease newsletter. 15(3). West Virginia Depart- 
ment of Agriculture. 

1982. 16(8). 

1983. Annual summary—1982. Cooperative Forest Pest Action Program. 14 








pp. 

BAKER, W. L. 1972. Eastern forest insects. USDA Forest Service Misc. Publ. No. 1175. 
642 pp. 

FORBES, W. T. M. 1948. Lepidoptera of New York and neighboring states, part II: 
Geometridae, Sphingidae, Notodontidae, Lymantriidae. Cornell Univ. Agric. Exp. 
Sta. Memoir 274. 263 pp. 

RINDGE, F. H. 1975. A revision of the New World Bistonini (Lepidoptera: Geometri- 
dae). Bull. Amer. Mus. Nat. Hist. 156:70-155. 

TALERICO, R. L. 1968. Life history of the looper Phigalia titea in Virginia. Ann. 
Entomol. Soc. Amer. 61:557-561. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 187-195 


THE RELATIONSHIP BETWEEN PEDALIODES PERPERNA 
AND PETRONIUS (SATYRIDAE), WITH THE 
DESCRIPTION OF A NEW SUBSPECIES 


LEE D. MILLER 


Allyn Museum of Entomology of the Florida State Museum, 
3621 Bay Shore Road, Sarasota, Florida 33580 


ABSTRACT. The species Pedaliodes perperna and P. petronius are compared and 
confirmed as separate species, rather than as forms of one species. Pedaliodes petronius 
kerrianna (La Mesa, near El Valle, Cocle, Panama) is described as new. The habits and 
habitat of the new subspecies are discussed and a possible foodplant association proposed. 
Adults and male and female genitalia of both species are illustrated and discussed. 


The interrelationship between Pedaliodes perperna (Hewitson) and 
P. petronius Grose-Smith has puzzled workers for years. Both species 
look rather similar, but there are significant differences in their sizes 
(petronius: forewing length about 35 mm, vs. less than 30 mm for 
perperna), wing shapes, maculation and the genitalia. Weymer (1912: 
253) classified petronius as a 2 form of perperna, the fact that the type 
of the former was actually a 6 notwithstanding. P. petronius is one of 
the largest Pedaliodes and certainly is the largest member of the genus 
occurring in Central America. The differences that are apparent when 
the two taxa are examined are of a nature that it seems incredible that 
they were ever considered to be conspecific. 

Superficial characteristics clearly separate petronius from perperna, 
but genitalic characters are even more dramatic. The penis of petro- 
nius (Fig. 14) is narrow, straight and not contorted in contradistinction 
to the contorted, stout penis of perperna (Fig. 13) and most other 
Pedaliodes. The é genitalia of P. petronius are somewhat reminiscent 
of those of Praepronophila Forster (see Forster, 1964:188, fig. 268), a 
very different butterfly in all other respects. The 2 genitalia are at least 
as dramatic, especially as regards the setose, ornate lamella antevagin- 
alis of petronius (Fig. 16), as opposed to the simple structure of per- 
perna (Fig. 15) and other, less spectacular differences. Perhaps petron- 
ius should be placed in another genus, but I have not yet examined 
enough “‘Pedaliodes’’ to determine whether this separation would be 
justified; therefore, petronius provisionally remains in Pedaliodes but 
as a species separate from perperna. 

Ecological separations also support the separation of the two species. 
P. petronius usually flies below 1000 m elevation wherever it has been 
recorded with adequate altitudinal data, whereas all of the specimens 
of perperna that I have seen have come from 1500-2000 m elevation. 
The ecological niche of petronius apparently is somewhat different, as 
is detailed in the description of the northern population of petronius. 


188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-4. Pedaliodes perperna (Hewitson). 1 & 2, syntype 6, upper (1) and under 
(2) surfaces, no data (Allyn Museum photo no. 040979-4/5; 3 & 4, syntype 9, upper (3) 
and under (4) surfaces, no data (Allyn Museum photo no. 040979-6/7). Both specimens 
are in British Museum (Natural History) collection. 


The differences here cited are certainly enough to ascertain that 
Pedaliodes perperna and P. petronius are separate species that replace 
one another altitudinally and are not even closely related within the 
genus Pedaliodes. 


Pedaliodes perperna (Hewitson) 
(Figs. 1-4, 18, 15a—b) 
Pronophila perperna Hewitson (1862:16-17). Type-locality not specified, but stated to 
be Venezuela by W. F. Kirby (1871:104). 6 and 2 syntypes in BM (see below) 
[examined]. 


Pronophila satyroides C. and R. Felder (1867:469-470). Type-locality: Caracas, Vene- 
zuela. Syntypes should be in BM, but not located. 


This species was described well by Hewitson (1862), and the ¢ gen- 
italia were figured by Forster (1964:166; fig. 224), though the orien- 
tation is different than that shown here. I can add little to the super- 


VOLUME 39, NUMBER 3 189 


ficial description of either the ¢ or of the 2, except to state that some 
Costa Rican 2 specimens have the extradiscal area of the upper fore- 
wing strongly laved with rufous, thereby setting off the ocellus in Cu,- 
Cu, more than is shown in the figure of a Venezuelan specimen. 

The ¢ genitalia (Fig. 13) are of the Pedaliodes type as illustrated by 
Forster (1964: figs. 198-260) with the contorted and complex penis. 
This organ is relatively shorter than is that of petronius, which in turn 
is relatively very straight and simple. The valva has a bilobate tip and 
no dorsal tooth like that which characterizes petronius. 

Female genitalia (Fig. 15) with simple, lightly sclerotized lamella 
ante- and postvaginalis, the latter with few setae. Width of lamella 
antevaginalis constant and attachment to ductus bursae simple. Antrum 
simple, ductus bursae only moderately sclerotized with paired dorsal 
supportive bars; ductus seminalis attached at juncture of ductus bursae 
and corpus bursae. Signa well developed with external spines. 

This species is apparently seldom common throughout its rather broad 
(for a Pedaliodes) range. I have seen specimens from Costa Rica, Pan- 
ama, Colombia and Venezuela, but from no locality have I enough 
material to pass judgment on possible geographic differentiation. All 
of the specimens have come from between 1500 and 2200 m elevation. 
By contrast, the following species is from a lower elevation, and two 
distinct geographic segregates have evolved. 

I had initially intended to designate the 6 specimen as the Lectotype 
for P. perperna, but several circumstances make this untenable. First, 
this specimen has no abdomen, hence its genitalia cannot be checked 
if the popular conception of perperna proves to encompass more than 
one species. Second, it is possible only by secondary sources to deter- 
mine from which population of the species the syntypes were taken. I 
suspect that Kirby (1871) was correct, and the specimens came from 
Venezuela, and further, it is likely that either the hills around Caracas 
or the vicinity of Colonia Tovar was the more precise type locality. 
This decision, however, should be left to some worker doing a com- 
prehensive revision of this group of Pedaliodes. 


Pedaliodes petronius petronius Grose-Smith 
(Figs. 5-8) 


Pedaliodes petronius Grose-Smith (1900:19). Type-locality: Valdivia, Colombia. HT in 
BM [examined]. 


Although this subspecies was described originally from Colombia, 
by far the majority of specimens have been from Panama, probably 
because of the recent emphasis on collecting in that country. Most of 
the specimens seen have come from the lower flanks of Panamanian 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 5-8. Pedaliodes petronius petronius Grose-Smith. 5 & 6, holotype 6, upper 
(5) and under (6) surfaces, COLOMBIA: ANTIOQUIA: “Valdevia” (=Valdivia) (Allyn 
Museum photo 040979-10/11), British Museum (Natural History) collection; 7 & 8, 9, 
upper (7) and under (8) surfaces, PANAMA: PANAMA; Cerro Jefe, 900 m, 9.iv.1977 
(Allyn Museum photo 090178-9/10), G. B. Small, Jr. collection. 


mountains (Cerro Jefe, Panama prov.), apparently in low montane 
forests (see below for details of the habitat of this species). 

The ¢ is totally fuscous above, and the 2? is only slightly irrorated 
with lighter scales toward the apex of the forewing. The under side is 
fuscous boldly marked postdiscally with tan on both wings and with a 
chestnut discal band; the ocelli stand out clearly against the ground 
color of both wings. 


VOLUME 39, NUMBER 3 191 





Fics. 9-12. Pedaliodes petronius kerrianna, new subspecies. 9 & 10, holotype 4, 
upper (9) and under (10) surfaces; PANAMA: COCLE: La Mesa, nr. El Valle, 820 m, 
5.11978; 11 & 12, paratype 2, upper (11) and under (12) surfaces, PANAMA: COCLE: 
La Mesa, nr. El Valle, 820 m, 5.i.1978. Both specimens are in Allyn Museum collection. 


é and 2 genitalia substantially the same as those of P. p. kerrianna 
(q.v., Figs. 14 and 16). 

It does not seem to be a common insect; I have seen only a few 
examples, but it may not always be rare where it is found if encoun- 
tered at the right time. Superficially, it seems to resemble members of 
the poesia group, but the genitalia are totally unlike any members of 
that complex. 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pedaliodes petronius kerrianna, new subspecies 
(Figs. 9-12, 14, 16a-—c) 


Male (Figs. 9-10). Head, thorax and abdomen dorsally covered with fuscous hairs; 
head and thorax with gray-brown ventral hairs, abdomen with buff ones ventrad. Palpi 
covered with fuscous dorsal and ventral and white lateral hairs. Legs with light gray- 
brown hairs. 

Upper surfaces of both wings deep fuscous with subapical buff patch from just outside 
forewing cell to 1A and usually small, barely noticeable black ocellus placed on inner 
part of buff patch in Cu,—Cu,. Wings with narrow dark marginal line following contours 
of wings. 

Under forewing basically fuscous with markings as in p. petronius, but differs as 
follows: between cell and margin is large buff patch speckled with fuscous scales, posi- 
tioned about as on upper side; black ocellus with white pupil in Cu,—Cu, larger than in 
p. petronius; subapical white ocellus in M,-M, much larger than that of petronius; and 
with supernumerary white subapical point in M,-M,. Under hindwing about as in p. 
petronius, except brighter and more contrasting and submarginal ocellus in Cu,—Cu, 
larger and more prominent. 

Male genitalia as illustrated (Fig. 14), similar to nominate subspecies, but dorsal tooth 
on valva somewhat less pointed and prominent. The long, straight penis and the toothed 
valva immediately distinguish this species from P. perperna. 

Length of forewing of holotype 6 34.1 mm; those of the 22 6 paratypes at hand range 
from 33.3 to 34.6 mm, averaging 34.14 mm. 

Female (Figs. 11-12). Head, thorax, abdomen and appendages about as in 6. Upper 
surface similar to nominate subspecies except for buff subapical forewing patch with its 
more prominent black, white pupilled ocellus in M,—M, of the same wing and at least 
hint of hindwing submarginal ocellus in Cu,—Cu,. 

Under surface much like that of p. petronius, but characterized by brighter buff 
forewing subapical patch, larger forewing and hindwing submarginal black ocelli with 
white pupils in Cu,-Cu, and generally more contrasting appearance. 

Female genitalia (Fig. 16) very ornate with numerous, presumably sensory scales and 
setae on VIII tergite and lamella postvaginalis. The normally membranous area anteriad 
of the papillae anales moderately sclerotized in this species; inner margin of lamella 
postvaginalis spinose and bearing two separate types of scales: a multidentate, short scale 
and a bidentate longer one. Lamella antevaginalis lightly to moderately sclerotized and 
cup-shaped; antrum heavily sclerotized and ornate. Ductus bursae similar to that of 
perperna, but tapered toward antrum. Signae longer than those of perperna. Attachment 
of ductus bursae to lamella antevaginalis (sterigma) much more heavily sclerotized than 
in perperna. 

Lengths of forewings of the four 2? paratypes at hand range from 36.0 to 37.4 mm, 
averaging 36.75 mm. 

Described from 59 specimens, 48 males and 11 females, from Cocle province, Panama. 

Holotype 6: PANAMA: COCLE: La Mesa, near El Valle, 820 m; 5.i.1978 (G. B. Small). 

Paratypes: 47 4, 11 2, same locality and collector as holotype, variously from November, 
December, January, June and July, 1978-1983. 

Disposition of type-series: holotype 6, 22 6 and four 2 paratypes in Allyn Museum of 
Entomology; 25 6 and seven 2 paratypes returned to Mr. Small for his collection and for 
distribution to other museums. 


This subspecies is named at Mr. Small’s request for Kerry Ann (Mrs. 
Robert) Dressler who discovered the original Panamanian colony. She, 
along with Mr. Small, has materially increased our understanding of 
Central American lepidopterology, especially of the fauna of Panama 
and Costa Rica. 

Mr. Small (pers. comm.) has written me extensively on the habits 


VOLUME 39, NUMBER 3 193 





1 mm 





Fics. 138 & 14. 4 genitalia of Pedaliodes. 13, P. perperna (Hewitson); genitalia prep- 
aration M-6653-v (Lee D. Miller); VENEZUELA: DIST. FED.: El Junquito. 14, P. 
petronius kerrianna new subspecies, paratype; genitalia preparation M-6599-v (Lee D. 
Miller); PANAMA: COCLE: La Mesa, nr. El Valle. Both specimens are in Allyn Museum 
collection. 


and habitat of this insect, as follows: “The locality is La Mesa, ca. 850 
m, above the town of El Valle de Anton, Cocle Province. The butterfly 
flies in a very wet thickety area with scattered thin trees festooned 
with moss and bromeliads (it does not fly in the surrounding thick 
forest). The average canopy height is about 15 ft., and paths through 
the area quickly become quagmires. Whether it is a natural formation 
or secondary succession following clearing of thick forest is problemati- 
cal. A thick-leaved Clusia (HYPERICACEAE) is abundant as are a 
number of MELASTOMACEAE, including Miconia oinochrophylla 
Donn.-Sm. and Tococca guianensis Aubl. Bromeliads on the ground 
are common, especially Guzmania musaica (Linden and Andre) Mez. 
in De. A broad-leaved cane type grass, Olyra standleyi Hitchc., is very 
abundant, and IJ strongly suspect that this is the larval food-plant. It is 
noteworthy that this grass is abundant near the top of Cerro Jefe, 
Panama Province, where typical petronius flies, and that this grass is 
not found in the thick forest, apparently needing a well lighted area 
in order to thrive.” 

Small states further, “In cloudy weather (which is most of the time) 
the butterfly makes rather short flights and alights low in the cane or 
other vegetation. However, when the sun shines brightly, it flies strong- 
ly and over the tops of the thickets and is very difficult to net. It 
generally flies from around 9:30 to 11:30 A.M. 

“TI have visited the locality in November, December, January, June 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 15 & 16. 9 genitalia of Pedaliodes species. 15, P. perperna (Hewitson); VEN- 
EZUELA: DIST. FED.: El Junquito; genitalia preparation M-6554 (Jacqueline Y. Miller); 
a: ventral view; b: dorsal view of sterigma and ductus bursae. 16, P. petronius kerrianna, 
new subspecies, PANAMA. COCLE: La Mesa, nr. El Valle; genitalia preparation M-6662 
(Jacqueline Y. Miller); a: ventral view; b: ventral view of sterigma and ductus bursae 
with lamella antevaginalis partially removed; c: dorsal view of sterigma and ductus 
bursae. 


and July and have observed or captured it on each occasion. It appears 
to be particularly numerous in November, although not abundant. At 
other times, it was in rather small numbers. It probably flies in every 
month, as I have records of typical petronius from Cerro Jefe in March 
and April.” 

An additional 2 specimen apparently of kerrianna was collected by 
Mr. Small at Moravia, Cartago, Costa Rica, at 910 m (3000 ft.) elevation 
on 27 July 1965. I have seen 18 other specimens collected by R. Hes- 
terberg at the same locality, though sometimes higher on the mountain. 
He states that petronius is found at elevations to nearly 1500 m, but 
that the ecological preference cited by Mr. Small is true of the Costa 
Rican specimens as well. These specimens and others have been ex- 
cluded from the type-series because the available material differs very 
slightly from the La Mesa specimens. P. J. deVries (pers. comm.) has 
reported other Costa Rican captures, and comments further on the 
ecological preferences of kerrianna. He says that it flies between 800 


VOLUME 39, NUMBER 3 195 


and 1000 m on the eastern slope of the Sierra de Talamanca, more 
specifically in the Valle de Reventazon along the Rio Pacadre in the 
Rio Chirripo region. He evidently has not found it at as high elevation 
as has Mr. Hesterberg. 


ACKNOWLEDGMENTS 


Many people provided information and inspiration for this article. Plant identifications 
were provided by Drs. Henry Stockwell and Annette Aiello of the Smithsonian Tropical 
Research Institute, Barro Colorado Island, Panama, and points of nomenclature (espe- 
cially the authors of the plants) were confirmed by Dr. David Hall of the Florida State 
Museum, Gainesville, Florida. The authorities at the British Museum (Natural History), 
London, England, especially Messrs. R. I. Vane-Wright and P. R. Ackery, allowed me 
access to the type specimens of both Pedaliodes petronius and perperna and greatly 
assisted me in other ways during my visit to London in 1979. Mr. Philip J. DeVries, 
Department of Zoology, University of Texas, Austin, Texas, and Richard L. Hesterberg, 
formerly of San Jose, Costa Rica, and now of Clearwater, Florida, provided habitat and 
range information on Costa Rican populations of kerrianna. Mrs. Robert (Kerry Ann) 
Dressler originally found the El Valle population of the butterfly named after her. The 
photographs were made by the late Dr. A. C. Allyn and my wife and colleague, Jacque- 
line, of this institution, and they also read and critically reviewed the manuscript. J. 
Miller also provided the female genitalic dissections and analyses. Mr. Michael J. Adams, 
Blandford, Dorset, England, answered innumerable questions about the taxonomy of 
Pronophilini. Clearly, Mr. Gordon B. Small, Jr., Balboa, Canal Zone, Panama, deserves 
special thanks, since it was he who first provided me with material of both petronius 
and kerrianna and gave the detailed habitat notes quoted herein. I must express my 
heartfelt gratitude to all of these people for their enthusiastic cooperation. 


LITERATURE CITED 


FELDER, C. & R. 1867 [1864-1867]. Reise der Osterreichischen Fregatte “Novara... 
Zool. 2. Lepidoptera. Carl Gerold’s Sohn, Wien. i-vi + 548 pp.; ill. 

FORSTER, W. 1964. Beitrage sur Kenntnis der Insektenfauna Boliviens XIX. Lepidop- 
tera III. Satyridae. Ver6ff. Zool. Staatsamml. Miinchen 8:51-188; ill. 

GROSE-SMITH, H. 1900. Description in Grose-Smith and W. F. Kirby, Rhopalocera 
Exotica. London, priv. publ., p. 19. 

HEwITsoN, W. C. 1862. On Pronophila, a genus of the Diurnal Lepidoptera; with 
figures of the new species, and reference to all those which have been previously 
figured or described. Trans. Entomol. Soc. London (3)1:1-17; ill. 

Kirby, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera. John Van Voorst, 
London. iii-v + 690 pp. 

WEYMER, G. 1912. Genus Pedaliodes, in A. Seitz, ed., Macrolepidoptera of the world 
5:250-262. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 196-200 


ECOLOGICAL NOTES ON SYNANTHEDON DOMINICKI 
DUCKWORTH AND EICHLIN (SESIIDAE) IN FLORIDA 
AND FIRST DESCRIPTION OF THE FEMALE 


LarRyY N. BROWN 
Department of Biology, University of South Florida, Tampa, Florida 33620 


THOMAS D. EICHLIN 


Insect Taxonomy Laboratory, A. & I., Division of Plant Industry, 
California Department of Food and Agriculture, Sacramento, California 95814 


AND 


J. WENDELL SNOW 
Fruit and Tree Nut Research Laboratory, U.S.D.A., Byron, Georgia 31008 


ABSTRACT. Synanthedon dominicki Duckworth and Eichlin, a clearwing moth 
previously known from only one specimen, was captured in sizeable numbers (40 spec- 
imens) in west-central Florida throughout most of the month of March 1985. The species 
responded only to the pheromone isomer (E,Z).2,13-octadecadienyl acetate and occurred 
only in cypress swamp habitat and the adjacent hydric hardwood forest ecotone. It was 
totally absent from nearby mesic and xeric plant communities. The first females of 
Synanthedon dominicki ever collected are also described. The species appears to be 
widely distributed in Florida but in a narrow ecological zone seldom collected for sesiids. 


The rare clearwing moth, Synanthedon dominicki Duckworth and 
Eichlin, is known from only one specimen taken on the Wedge Plan- 
tation, South Santee River, Charleston County, South Carolina on 27 
March 1967 (Duckworth & Eichlin, 1973). The holotype is a male 
collected at a black light, and the female until now was unknown. 

Throughout the early months of 1985, several sesiid pheromones, or, 
more correctly sex attractants, were employed to survey clearwing 
species at numerous locations in west-central Florida. The initial chem- 
ical isomer of the attractants was identified by Tumlinson et al. (1974), 
and the isomer effective for attracting S. dominicki was first identified 
by Schwarz et al. (1983). Several permanent sampling stations using 
sticky traps were placed in natural plant communities located on the 
500 acre Ecological Research Area of the University of South Florida 
in Hillsborough County. Beginning on 4 March 1985, male S. dom- 
inicki began appearing in traps at two of the sampling stations. One 
group of traps was located in the center of a bald cypress (Taxodium 
distichum) swamp, and the other group was situated in a stand of 
water oaks (Quercus nigra) adjacent to the cypress swamp. A total of 
40 male S. dominicki were taken in several traps baited with the at- 
tractant (E,Z) 2,13-octadecadienyl acetate between 4 March and 31 
March. Most of these males (25) were trapped during a ten day period 


VOLUME 39, NUMBER 3 197 





Fics. 1 & 2. Adults of Synanthedon dominicki: 1, male; 2, female. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


198 





< 


YY) 
ASS 







We 


vi 


\ 


ie 





Synanthedon dominicki: 3, female genitalia (ventral view); 4, male 


Fics. 3 & 4. 
genitalia (ventral view, left valve removed). 


VOLUME 39, NUMBER 3 199 


in mid-March. S. dominicki failed to respond to any of five other 
isomers used, and none was trapped in either February or April in the 
Tampa Bay area, even though all traps were baited continuously dur- 
ing those months also. Thus, the emergence window for adults was 
rather short and confined to early spring. This species appears to be 
quite habitat specific, because no males were taken in numerous traps 
placed in plant communities located just outside the swamp ecotone, 
such as pine flatwoods, pine-turkey oak, oak-palmetto scrub, live oak 
hammock, wax myrtle-brush, or old-field communities. 

Traps were checked at hourly intervals for several days in mid- 
March to determine the duration of the daily flight period for male S. 
dominicki. They entered traps only between 1300-1600 h, with the 
greatest flight activity occurring at mid-afternoon. 

Two additional male S. dominicki were taken in sticky traps set 5- 
14 April 1985 in a swampy hardwood forest near Crystal River, Citrus 
County, Florida. This location is approximately 90 mi. north of the 
permanent sampling stations in the Tampa Bay area. Since this record 
is at least a week later than the end of the flight observed this year 
near Tampa, it suggests that the emergence period for the moth may 
be sightly later in north-central Florida or last somewhat longer. 

While checking traps at the border of the cypress swamp on 10 
March 1985, a single female S. dominicki was hand-netted as she hov- 
ered in the vicinity of a small waterlocust (Gleditsia aquatica). She 
was captured about 1600 h, while observed to be intermittently landing 
on and hovering above the vegetation of this tree. The host plant of S. 
dominicki is unknown, but it could not be determined if she was laying 
eggs on the waterlocust. This specimen constitutes the first record of 
the female of this rare clearwing moth, and the description follows 
below. 

Surprisingly, a second female was collected in an insect flight trap 
on 15 April 1985 in a swampy area near White Springs in Columbia 
County, Florida. This is a northern Florida location about 200 mi. north 
of the Tampa Bay area and fits the postulated later emergence window 
as one proceeds northward. 


Description of Synanthedon dominicki 


Female (Fig. 2): Head. Front blue-black, some white adjacent to eyes; vertex blue- 
black, orange mixed posteriorly; occipital fringe orange; labial palp smoothly scaled, 
orange with black apically; antenna blue-black, lacking ventral cilia and lacking preapical 
white spot. 

Thorax. Blue-black, narrow subdorsal stripe, orange beneath wings; legs blue-black, 
white on spurs and at joints of tarsal segments. 

Wings. Forewing length: 9-10 mm. Forewing opaque, blue-black, ventrally with yel- 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


low at wing bases; hindwing hyaline with brown-black on small, narrowly triangular 
discal spot and with some diffuse dark scaling apically. 

Abdomen. Entirely blue-black except for orange-red on posterior end, including anal 
tuft and tip of abdomen. 

Female genitalia. Structures as shown (Fig. 3), with well-differentiated ostial region, 
and membranous narrow pouch-like protrusion on corpus bursae. 

Male (Fig. 1) (see female description above and original description of species, Duck- 
worth & Ejichlin, 1973): Specimens differ slightly from male holotype in that the antennae 
lack pre-apical white spots, and the vertex of the head has orange mixed at the posterior 
margin. The major difference from the female is the forewing opaqueness, males with 
basal one-half hyaline, females entirely opaque. 

Forewing length of males. 7-9 mm. 

Male genitalia. As illustrated (Fig. 4). 

Host plant. Unknown. 

Distribution. Using sex attractants, the range has now been extended from South 
Carolina (type locality) to Georgia, Alabama and Florida. 

Discussion. The data for the first recorded female specimen is as follows: Ecological 
Research Area, University of South Florida, Tampa, Hillsborough County, Florida, 10 
March 1985, collected by L. N. Brown. This specimen will be deposited in the collection 
at the U.S. National Museum of Natural History, Washington, D.C. 


ACKNOWLEDGMENTS 


We thank Kathy Scarborough, Research Technician, USDA Lab. Byron, Georgia for 
helping to prepare and deploy traps in Georgia and Alabama, preparation of specimens 
and other forms of technical help throughout this and other ongoing clearwing moth 
studies. Thanks also to Charles S. Papp, Sierra Graphs and Typography, Sacramento, 
California for applying the final inking to the illustrations and photographing the adult 
moths. 


LITERATURE CITED 


DuckwortTH, W. D. & T. D. EICHLIN. 1973. New species of clearwing moths (Lepi- 
doptera: Sesiidae) from North America. Proc. Entomol. Soc. Wash. 75:150-159. 
SCHWARZ, M., J. A. KLUN, B. A. LEONHARDT & D. T. JOHNSON. 1983. (E,Z)-2, 13- 
octadecadien-1-ol acetate. A new pheromone structure for sesiid moths. Tetrahedron 
Letters 24:1007-1010. 

TUMLINSON, J. H., C. E. YONCE, R. E. DOOLITTLE, R. R. HEATH, C. R. GENTRY & E. 
R. MITCHELL. 1974. Sex pheromones and reproductive isolation of the lesser peach- 
tree borer and peachtree borer. Science 185:614-616. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 201-207 


HOW TO DO GENETICS WITHOUT MAKING 
THE BUTTERFLIES CROSS 


JOHN R. G. TURNER 


Department of Genetics, University of Leeds, 
Leeds LS2 9JT, England 


ABSTRACT. It is possible to find out whether an inherited variety of a butterfly is 
sex-linked and, if it is not sex-linked, whether the variety is dominant or recessive to 
normal (in short, to find out its basic genetics) without carrying out pedigreed breeding 
experiments. These require much space, time and record-keeping, and are in any case 
not possible in some species. 

Instead, one can raise offspring from the two types of female captured in the wild or 
followed while ovipositing. The mates of the females need not be observed. A fairly 
simple calculation based on the numbers of the two types of offspring produced by the 
two types of female will then reveal the inheritance of the variety. 

The method is illustrated with data on the green and yellow forms of the African 
Papilio phorcas. 


Working out how the different forms of a butterfly are inherited 
can be tedious; a pedigree record must be kept over a number of 
generations, the offspring of different females must be kept separate, 
and one needs to be fairly skilled in mendelism to set the crosses up 
in the way that will give the necessary information. If the variant one 
is studying is confined to the female, as are the white forms of some 
Colias or the black form of Papilio glaucus, the exercise becomes even 
harder, for as one cannot tell what color the male “ought” to be, one 
must do the crosses “blind.” In addition, some butterflies cannot easily 
be mated in captivity. 

It is, however, possible to do butterfly genetics without any of this 
hassle. Provided wild caught females can be persuaded to lay eggs or 
can be found ovipositing in the field, it is possible to determine the 
genetics of naturally occurring forms simply by raising the offspring 
of wild females. Neither the possibility that the female may be pro- 
ducing a mixed brood after mating with two males, nor even combin- 
ing the offspring of different females in one breeding cage, will spoil 
the method. The only requirements are that one must be certain which 
color of female was the mother of the eggs, and that one of the forms 
should be rarer than the other. (When the forms are exactly equal in 
frequency the method fails completely and it requires very large num- 
bers of offspring indeed when the rare form is over around 35% of the 
population.) With some tropical species one must be very cautious 
about information obtained from whole egg rafts, as these are some- 
times laid cooperatively by several females; unless all the females have 
been seen from the laying of the first egg, and all are of the same 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


color, the result can be completely unreliable (Turner, 1971, 1981; 
Mallett & Jackson, 1980). 

I will describe the method for a butterfly having just two forms, and 
use data on the green and yellow forms of the African Papilio phorcas 
for illustration. North temperate zone workers may find it easier to 
think in terms of Colias: to do this, simply substitute mentally “white’’ 
for “green.” With three or more forms the method becomes, needless 
to say, more complicated. 

The method depends on a principle readily derived from the tenets 
of population genetics, that if the females of a rare form, having mated 
randomly with the males in their population, give rise to offspring 
which are mostly of the common form, then the rare form is recessive. 
On the other hand, if the rare form is dominant, it will give this fact 
away by producing among its offspring roughly equal numbers of the 
two forms. The common form, whether dominant or recessive, always 
tends to produce a majority of offspring like itself. 

Some mathematical precision can be given to this idea (I give the 
proof elsewhere—Clarke et al., 1985). If the frequencies of the domi- 
nant and the recessive genes (not forms) in the population are p and 
q, then the recessive females, overall, produce offspring in just these 
proportions. If the green form of Papilio phorcas was recessive and 
the gene frequencies for yellow and green were 75% and 25%, then in 
aggregate a sample of eggs from a number of green females would 
produce 75% yellow and 25% green offspring. So, calling D the pro- 
portion of dominants in the offspring and R the proportion of reces- 
sives, we have the formula for the offspring of recessive females: 


D = p, R=G (1) 


where p is the frequency of the dominant gene and q the frequency 
of the recessive. 

Dominant females on the other hand produce the two forms ac- 
cording to the formula 


iE) 
R 


1 
aaa @ 


where p and q are as before the frequencies of the dominant and 
recessive genes. 


If the gene frequencies were as before, but the green form was 
dominant, then in the offspring of green females we would have 


D/R = 0.25/0.75 + 1/0.75? = 2.11 


and 


VOLUME 39, NUMBER 3 203 


TABLE 1. Offspring of wild females of Papilio phorcas from Nairobi and Ngong 
(Kenya). From Clarke et al. (1985). 


Mother 
Offspring Green Yellow 
Green 78 (84%) 37 (51%) 
Yellow 15 (16%) 35 (49%) 


D = 2.11/(1 + 2.11) =068, R=1/(1 + 2.11) =0.32 


so that 68% of the offspring of green females would be green and 32% 
yellow. 

To determine whether a rare form is dominant or recessive, we 
therefore compare the frequencies of the two forms among its offspring 
with those we would expect according to formula (1) and formula (2). 
Provided the butterflies are mating at random, one of these formulae 
will give an answer fitting the data, and the other will not. Obviously, 
to do this we need to know the values of p and q which, as we will 
see, can be obtained either from a population sample or from further 
breeding work. 

What is needed, therefore, is a set of offspring derived from females 
of the rare form. There is no need for the mate of the mother to be 
known, nor to have any minimum number of offspring from any one 
female (they could well be eggs found by following ovipositing females 
around in the field), nor any need to keep the offspring of different 
females separate. All that is needed is the certainty that they are the 
offspring of the rare type of female. 

In addition, it is necessary to have an estimate of the frequency with 
which the rare form occurs in the population, obtained by catching as 
many individuals as possible without making a special effort to capture 
either kind, or provided the population is large and the butterflies not 
too sedentary, simply by keeping a tally of the numbers of the two 
forms seen. If this is not obtainable, a satisfactory substitute is a large 
set of offspring derived from the commoner kind of female. Again, so 
long as they certainly are from this type of female, no further infor- 
mation is needed. 

In sum, we need (1) a set of offspring from the rarer type of female, 
plus (2) either a field estimate of the proportions of the two forms or 
a set of offspring from the commoner type of female. Data of this kind 
for P. phorcas are shown in Table 1, where I have combined all the 
offspring of a large number of wild green mothers and of yellow wild 
mothers from the Nairobi area (including the town of Ngong). In ad- 
dition, the yellow form has been reported as rare in this region, prob- 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ably being a little less than 20% of the population. Suppose first that 
yellow is recessive. The frequency of the gene is then given by 


q = V0.2 = 0.447 


(a surprisingly large frequency—nearly 45%—as recessive genes are 
always much more common than the form which they control). As the 
yellow form is recessive, yellow females should produce yellow and 
green offspring, from formula (1), in the proportions 


yellow = q = 0.447 
green = p= 1 — q= 0.558 


These proportions are close to the observed numbers of the two kinds 
of offspring and we strongly suspect that the yellow form is recessive. 

Does the hypothesis that the yellow form is dominant fare worse? 
In that case the frequency of the green gene (which must be recessive) 
is 


qg= V1 — 0.2) = 0.894 


As the yellow form is dominant, it will give rise to yellow and green 
forms, according to formula (2), in the ratio 


yellow/green = p/q + 1/q? 


or in this case 1.368 : 1. This means that among the offspring we expect 


1.368/(1 + 1.868) = 0.578 yellow and 
1/(1 + 1.368) = 0.422 green 


which is not such a good fit to what is actually observed (Table 1). The 
yellow form therefore appears to be recessive. 

However, suppose that we do not have a good estimate of the fre- 
quencies of the two forms in the population (and the estimate of 20% 
yellow is in fact not particularly accurate). A perfectly good substitute 
for this estimate is the number of the two forms appearing among the 
offspring of the common female form. Our data for the numbers of 
yellow and green females arising from green mothers are given also in 
Table 1. 

Start by supposing that yellow is dominant. In that case the yellow 
and green proportions from the green mothers are direct estimates of 
the gene frequencies p and q, giving in this case p = 0.161 (yellow) 
and 0.839 (green). We can test this against the offspring of yellow 
females, again by using the formula 


yellow/green = p/q + 1/q? 


VOLUME 39, NUMBER 3 205 


and in this case yellow: green is 1.614:1; yellow individuals should be 
1.614/(1 + 1.614) = 0.617 and green individuals 1/(1 + 1.614) = 0.383 
of the offspring. Again, the fit is not very good. 

Checking whether yellow being recessive gives a good fit is harder 
this time. If yellow is recessive the ratio of green/yellow from green 
mothers, which from Table 1 can be calculated as 78/15 = 5.2, will 
give q if we solve the equation 


(UO) Gt WGP 


This is a quadratic in q, and according to standard algebra, the general 
solution is that if x is the ratio of green to yellow from green mothers, 
then 


Jb ae W483 SPS) 


ape 9 (3) 


q= 
Substituting 5.2 for x in (8) gives us q = 0.490 and therefore p = 0.510. 
These should be the proportions of yellow and green among the off- 
spring of yellow mothers, which is clearly an excellent fit (Table 1). 
The yellow form is obviously recessive. 

The results, particularly if numbers are small, might not be so ob- 
vious as this, and then a statistical test would have to be applied, 
comparing the observed and expected numbers (not the percentages). 

We can summarize the value of the method with Table 2. The first 
column shows the frequency of the form which is actually recessive, 
and the next the frequency of the recessive gene. If we obtained off- 
spring from recessive females (which are the rare form above the line 
and the commoner form below it) we would obtain the offspring pro- 
portions shown in the third column; the fourth column shows the off- 
spring which would be obtained from the dominant females (which 
are the rare form in the lower half of the table). The last column shows 
the proportions which we would calculate for the offspring of rare 
females (recessive above the line, dominant below) when we took the 
wrong hypothesis about the dominance. By comparing this with the 
numbers in bold type, we can see how easy, or not, it is to tell that we 
are in fact wrong. It can be seen that provided one or other form is 
below about 30%, the method will distinguish very well which of the 
forms is dominant but that it will not work when the forms are nearly 
equally common in the population. 

It is, however, still worth making the observations even when the 
forms are equally abundant, for this allows us to distinguish a sex- 
linked gene. For if the gene were carried on the X chromosome, then 
both kinds of female would produce offspring in the same proportions: 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Proportions of two types of offspring from both types of mother at different 
population frequencies of the rare form. 


Wrong assumption about 
Actual frequency Actual frequency Recessive (yellow) females Dominant (green) females dominance of rare form 
of recessive of recessive give dominant: recessive give dominant:recessive _ predicts that it will give 


form (yellow) gene (yellow) (green: yellow) (green: yellow) green: yellow 
0.001 0.032 0.968:0.032 0.999:0.001 0.499:0.501 
0.005 0.071 0.929:0.071 0.995:0.005 0.496:0.504 
0.01 0.100 0.900:0.100 0.991:0.009 0.493:0.507 
0.05 0.224 0.776:0.224 0.959:0.041 0.4:70:0.530 
0.1 0.316 0.684:0.316 0.924:0.076 0.444:0.556 
0.2 0.447 0.553:0.447 0.862:0.138 0.399:0.600 
0.3 0.548 0.452:0.548 0.806:0.194 0.360:0.640 
0.4 0.633 0.367:0.633 0.755:0.245 0.325:0.675 
0.5 0.707 0.293:0.707 0.707:0.293 0.293:0.707 
0.6 0.775 0.225:0.775 0.662:0.338 0.633:0.367 
0.7 0.837 0.163:0.837 0.619:0.381 0.548:0.452 
0.8 0.894 0.106:0.894 0.578:0.422 0.447:0.553 
0.9 0.949 0.051:0.949 0.538:0.462 0.316:0.684 
0.95 0.975 0.025:0.975 0.519:0.481 0.224:0.776 
0.99 0.995 0.005:0.995 0.504:0.496 0.100:0.900 
0.995 0.998 0.002:0.998 0.502:0.408 0.071:0.929 
0.999 0.9995 0.0005:0.9995 0.5004:0.4996 0.032:0.968 


The ease with which one can tell which form is in fact recessive can be seen by comparing, in any particular row, 
the figures printed in bold type. Within the dotted lines, the figures are well-matched and the dominance is hard to 
determine; above and below these lines there is clear discrimination, and this is particularly marked when the recessive 
form is very rare or very common, as at the top and bottom of the Table. 


say 60:40 green and yellow from both green and yellow mothers. 
Whereas, if the gene is not on the sex chromosome, the proportions, as 
can be seen from the center line of Table 2, are mirror images; the 
yellow form produces yellow: green in the ratio 0.71:0.29, whereas 
green produces them in the ratio 0.29:0.71. 

It should be noted that this method becomes completely unreliable 
if the offspring of pedigreed captive matings are included in the data; 
the only permissible use of captive bred butterflies is to take virgin 
females and mate them to wild-caught males, or to collect larvae at 
random in the wild and then test their offspring, for the first generation 
only, by mating them in captivity. Indeed, when I first tried to apply 
the method to Papilio phorcas, there were few matings and I included 
the offspring of some pedigree broods to swell the numbers; the method 
then gave the totally incorrect answer that yellow was dominant, which 
shows how unreliable it is in those circumstances. 

I believe that useful information could be obtained on the genetics 
of some of the more “difficult’’ species of butterflies and moths, by 
using this technique. The recipe provided above will be found quite 
easy to follow if it is applied step by step. As an example, readers 


VOLUME 39, NUMBER 3 207 


might like to try to determine the inheritance of an imaginary white 
Colias, occupying 3% of its population, and giving 51 white to 49 
orange from white mothers. 

As a matter of history, it is worth recording that the first use of a 
primitive version of this method appears to have been by E. B. Poulton 
(1914), who determined in this way that one of the rare forms of 
Papilio dardanus was produced by a dominant gene. 


ACKNOWLEDGMENTS 


I am most grateful to Professor Sir Cyril Clarke, KBE, FRS and to Lady Clarke, who 
suggested this problem and the broad outline of its solution to me and who provided the 
data on Papilio phorcas. They also read the draft and suggested some improvements. 


LITERATURE CITED 


CLARKE, C. A., F. M. M. CLARKE, S. C. COLLINS, A. C. L. GILL & J. R. G. TURNER. 
1985. Male-like females, mimicry and transvestism in butterflies (Lepidoptera: Pap- 
ilionidae). System. Entomol. 10:257-288. 

MALLET, J. L. B. & D. A. JACKSON. 1980. The ecology and social behaviour of the 
neotropical butterfly Heliconius xanthocles Bates in Colombia. Zool. J. Linn. Soc. 
London 70:1-18. 

POULTON, E. B. 1914. The Mendelian relationship of the female forms of P. dardanus. 
Proc. Entomol. Soc. London 1914, Ixvii-lxx. 

TURNER, J. R. G. 1971. Studies of mtillerian mimicry and its evolution in burnet moths 
and heliconid butterflies. In E. R. Creed (ed.) Ecological genetics and evolution, pp. 
224-260. Oxford, Blackwell. 

1981. Evolution and adaptation in Heliconius: a defence of neo-Darwinism. 

Ann. Rev. Ecol. Syst. 12:99-121. 





Journal of the Lepidopterists’ Society 
39(3), 1985, 208-214 


NOTES ON PSEUDOSPHINX TETRIO (L.) 
(SPHINGIDAE) IN PUERTO RICO 


JORGE A. SANTIAGO-BLAY’ 


Biology Museum, Biology Department, University of Puerto Rico, 
Rio Piedras, Puerto Rico 00931 


ABSTRACT. An egg cluster of Pseudosphinx tetrio (L.) (Lepidoptera:Sphingidae) 
was reared to determine the duration of each developmental stage. The adult emerges 
after 53 days following oviposition: eggs eclose in three or more days (n = 90, sd = 0); 
the mean time span of the five larval instars is 24 days (n = 22, sd = 0.8), or if six stages, 
29-30 days (n = 2); prepupa, close to four days (n = 22, x = 3.8, sd = 0.5); pupa, about 
22 days (n = 22, x = 22.2, sd = 0.5). Larvae feed on Apocynaceae such as: Plumeria 
spp., Allamanda cathartica, and A. violacea. All stages, including the egg, are illustrated 
and briefly described. The morphometric variation of most stages is reported, as well as 
notes on the coloration of the newly molted larvae and pupae, and other data on the 
biology of the species. 


One of the most popular ornamental trees in Puerto Rico is Plumeria 
rubra L. (Magnoliophyta: Apocynaceae), known locally as ramo or 
pucha de novia, frangipani or aleli. Especially during July to Septem- 
ber, P. rubra trees are attacked by larvae of the sphingid moth Pseu- 
dosphinx tetrio (L.) (Fig. 2), which can defoliate and deflower a tree 
in a few days. Pseudosphinx tetrio has been reported as feeding in 
other Plumeria species in Puerto Rico such as P. alba and P. obtusa 
(Martorell, 1976) but has not been recorded feeding on species in other 
genera. The association of P. tetrio with Plumeria spp. was suggested 
by Fabricius (1775), when he described the moth under the name 
Sphinx plumeriae (Cadiou, pers. comm.). Haber (1984) described the 
floral biology of P. rubra in Costa Rica. 

This moth is widespread throughout the American tropics and has 
been reported from the southern United States (McDonnough, 1938; 
Hodges, 1971) to Paraguay and southern Brazil (Moss, 1920; Forbes, 
1930). In Central America it has been suggested as a possible coral 
snake mimic (Janzen, 1980). This species is also known from the Ca- 
ribbean, having been reported from Cuba under the name of Sphinx 
asdrubal (Poey, 1832), Jamaica (Gundlach, 1891), the Dominican Re- 
public (Druce, 1881-1900), and the Puerto Rico Region (Dewitz, 1877; 
Gundlach, 1891; Forbes, 1930; Martorell, 1945, 1976; Wolcott, 1948; 
and Medina-Gaud & Martorell, 1974). 

There is only one nearly complete account of the duration of de- 
velopment of this species (Dinther, 1956), although partial observations 
have been reported earlier (Merian, 1726; Sepp, 1852, both cited by 
Dinther; and Janzen, 1983). 


' Present postal address: Department of Entomological Sciences, University of California, Berkeley, California 94720. 


VOLUME 39, NUMBER 3 209 


The purpose of this paper is to report the duration of the develop- 
mental stages in the life history of P. tetrio. I will also add other 
biological information that was gathered during the study. 


MATERIALS AND METHODS 


An egg cluster was collected during the afternoon of 14 August 1982 
in a xerophytic forest near road 333, 8 km from Guanica, a small town 
in southwest Puerto Rico. The cluster was placed in a polyurethane 
box, subsequently transferred to a plastic jar, and then placed in an 
incubator at 24—-26°C, 24 h darkness. Humidity was provided by plac- 
ing wet pieces of towels in the jars. After eclosion 30 larvae were placed 
individually in 30 ml cups and fed with P. rubra (red variety). After 
the third instar each larva was transferred to a 1000 ml jar. The con- 
tainers and the larvae were cleaned with tap water at least every two 
or three days. When the pupal stage was reached, all food was removed 
from the jars. Prior to the emergence of adults, a piece of the central 
vein of a P. rubra leaf was placed in the jar in order to facilitate 
climbing and wing expansion. After emergence, several adults of each 
sex were kept in a 0.1 m? plastic box and fed with sugar water. 

A previous partial rearing, from which data was taken on prepupal 
and pupal weight loss and some of the stage duration, had been done 
under similar conditions except for the dark-light period which had 
been about 12-12 h. 

Eggs were measured with an ocular micrometer. All other measure- 
ments were made, usually during the first day of the appearance of 
the instar, using a metric ruler except for the cephalic width which 
was measured with a caliper. To measure the length of the larvae, the 
insect was allowed to remain inactive, and then it was firmly held by 
the extremities and measured. 

Sixty other larvae, kept in groups of less than 12 specimens, were 
regularly fed with leaves of the apocynaceans Allamanda cathartica 
L., A. violacea G. Gardn. & Fielding, Neriuwm oleander L., and occa- 
sionally, P. rubra. 


RESULTS AND DISCUSSION 


Egg 

The egg cluster consisted of 96 eggs (Dinther, 1956, reported a max- 
imum of 69) and was found on the upper surface (Dinther, 1956, 
reported clusters only on the lower surface) of a Plumeria obtusa leaf. 
All but two eggs were laid in a single layer and generally, very close 
to or touching each other (Fig. 1). The cluster lacked a cover like that 
secreted by other moths. The eggs showed no sculpturing except for 
minute punctures on the surface. They were pale green, ellipsoidal, 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1 & 2. Pseudosphinx tetrio stages: 1, egg cluster; 2, fifth larval instar. 


and measured 2.2 x 2.5 mm (n = 11, sd < 0.1). Ninety of the 96 eggs 
hatched, two larvae were found with part of their intestine protruding 
from a lateral injury, which I presumed was caused either by the edges 
of the shell or by cannibalism of other larvae. Two of the six eggs that 
failed to hatch had a small hole in one of the extremes (parasitism?). 
Eclosion takes place synchronously, at least three days (n = 90, sd = 
0) after oviposition. 


Larva 


The color of the newly molted, eruciform larva of each instar is light 
yellow and dark gray in alternating transverse rings. Several hours 
later, the larvae acquired the typical yellow and black coloration (Fig. 
2). 

Larval length and head capsule width increase in successive instars 
but tail length decreases after the third instar, probably due to reab- 
sorption or to breakage followed by the production of a new but shorter 
tail. Table 1 summarizes the morphometric variation among these 
structures throughout the five, or six, larval instars. The results are in 
general agreement with those reported by Dinther (1956). 


VOLUME 39, NUMBER 3 211 


TABLE 1. Morphometric variation of total larval length, cephalic capsule width, and 
tail length in Pseudosphinx tetrio (L.). All measurements in mm. Usually there are five 
larval instars, a sixth was reached by a few specimens, two of which did not pupate. 


n Instar 
Character x pe EE re ee ae 
sd 1 2 3 4 5 (6) 


Total larval length 30 29 29 28 28 4 
17.0 13.0 20.7 34.3 63.0 69.3 
0.4 1.9 2.8 4.0 9.4 5.7 

Cephalic capsule width 30 29 282 28 28 4 
1.0 2.0 3.0 4.8 Tel 8.8 
0 0.2 0 0.4 0.7 0.5 

Tail length 30 29 2. 28 rat 3? 
3.9 7.3 12.4 11.8 Med 9.7 
0.3 Jig) Lee 3.1 4.6 5.0 


* One measure not taken. 
> Two broken tails, not measured. 


Larvae fed with P. rubra passed through five larval instars, less 
frequently six. Fifth instar larvae can consume three to four leaves per 
day. The mean duration of each instar is: lst = 3.2 days (n = 29, sd = 
0.5); 2nd = 4.2 days (n = 29, sd = 0.5); 8rd = 4.6 days (n = 28, sd = 
0.9); 4th = 5.5 days (n = 28, sd = 0.8); 5th = 6.5 days (n = 24, sd = 
1.0); a mean total of 24.0 days (n = 22, sd = 0.8). If a sixth instar is 
present, eight more days are needed for a total of 29-30 days. Larvae 
that reach a sixth instar have a shorter fourth and fifth instars (4.5 and 
5 days, respectively). 

Other larvae were fed leaves of Allamanda cathartica and Nerium 
oleander, after being fed during the first two days with P. rubra leaves. 
Offered alone, A. cathartica leaves were eaten slowly, but those of N. 
oleander remained almost untouched. Other larvae were fed with A. 
violacea leaves. 

Based on the available information, I presume that P. tetrio larvae 
rarely feed on Allamanda spp. in nature. An examination of the col- 
lection and of the accession cards at the Entomology Museum, Agri- 
cultural Experiment Station, Rio Piedras, Puerto Rico, revealed only 
one record of P. tetrio larvae feeding naturally on A. cathartica. Al- 
lamanda cathartica constitute a new plant host record for P. tetrio in 
Puerto Rico. Other larvae were offered leaves of Pterocarpus indicus 
(Fabaceae), Carica papaya (Caricaceae), Bambusa vulgaris (Poaceae), 
Lagerstroemia speciosa (Lythraceae), Calotropis procera (Asclepia- 
daceae), and Wedelia trilobata (Asteraceae), but all the leaves re- 
mained untouched and many larvae died of starvation. 


212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 3-5. P. tetrio stages: 3, pupa; 4, male; 5, female. 


Prepupa 


This stage is characterized by the shortening and darkening of the 
body, reduction of the prolegs, and bending of the tail. By the end of 
the prepupal stage, the larva has spun a silken case using also part of 
the uneaten foliar material. This period lasts almost four days (n = 22; 
xk = 3.8, sd = 0.4; n = 30; x = 4.0; sd = 0.2 in the previous partial 
rearing). During the prepupal stage the organism loses 38.9% of its 
weight at the beginning of this stage (initial mean weight = 12.15 g; 
sd = 2.64; final mean weight = 8.74 g; sd = 1.68; n = 24). 


Pupa 


The newly formed pupa is yellow. After the second to the third 
hours, brown spots appear on its surface, and by the sixth hour the 
color has darkened to yellowish brown with lateral dark stripes on the 
thorax and rings on the abdomen. Later the pupa acquires the typical 


VOLUME 39, NUMBER 3 213 


uniformly dark brown coloration (Fig. 3). During this stage there is a 
mean weight loss of 18.2% (initial mean weight = 8.74 g, sd = 1.68; 
final mean weight = 7.15 g, sd = 1.45; n = 24). Female pupae are 
slightly longer (L) and wider (W) than males (2, n = 10, x, = 72.1 mm, 
seep Xo — 17.2, sdy — 1.1; ratio L/W = 4.2; 6, n = 12, x, = 69.6, 
sd, = 2.0; xy = 16.0, sdy = 0.4; ratio L/W = 4.4). The pupal stage 
lasts about 22 days (n = 22, x = 22.2, sd = 0.5; n = 80, x = 21.4, sd = 
0.6 for a previous partial rearing). 


Adult 
(Figs. 4, 5) 


Adult females measure slightly more in length (L), width (W), and 
wingspan than males (2, n = 7, x, = 59.7 mm, sd = 2.8; Xyw = 15.8, 
sdy = 1.2; n = 3, wingspan range 134-150; 6, n = 9, x, = 56.7, sd, = 
4.4; xy = 13.9, sd, = 0.6; n = 4, wingspan range 107-129). Wingspan 
values are similar to those reported by Dinther (1956). Adults kept in 
captivity lived up to 10 days but no eggs were laid. 

It is interesting to speculate about why this cycle remained little 
known for such a long time. Apparently, this is partly due to the 
susceptibility of some stages to infections (Moss, 1920; Janzen, 1983; 
and Abreu, pers. comm.). In addition, earlier reports lack information 
about rearing conditions, which may have been inappropriate and might 
have caused unsuccessful rearing attempts. 


ACKNOWLEDGMENTS 


I wish to thank my wife, Maria Esther Arroyo-Sanchez, for her collaboration in dif- 
ferent stages of the laboratory work. Mr. Vincent Lee (Cal. Acad. Sci., San Francisco, 
CA) and Mr. William L. Murphy (IIBIII, Beltsville, MD) provided photocopies of some 
‘papers that were not available to me. Dr. José A. Mari Mutt (Univ. P.R., Biology De- 
partment, Mayagiiez) and Dr. Daniel H. Janzen (Department of Biology, Univ. Penn- 
sylvania, Philadelphia, PA; and Parque Nacional Santa Rosa, Liberia, Costa Rica) read 
the manuscript and suggested valuable changes. Mr. Rafael Ingles (Crop Protection Dept., 
Univ. P.R., Mayagiiez) kindly provided me with the record of Pseudosphinx tetrio feed- 
ing on Allamanda cathartica in nature. Mr. Edwin Abreu (Crop Protection Department) 
provided the information on the susceptibility of P. tetrio to infections. Dr. Jean-Marie 
Cadiou pointed cut the Fabricius reference. 


LITERATURE CITED 


DewitTz, H. 1877. Damerungs- und Nachfalter von Porto Rico, gessalmmelt von Herrn 
Consul Krug. Mitt. Munch. Ent. Ver. 1:91-96. 

DINTHER, J. B. M. 1956. Three noxious Hornworms in Suriname. Entomol. Ber. 16: 
12-15. 

Druce, H. 1881-1900. Insecta. Lepidoptera—Heterocera. In Biologia Centrali-Ameri- 
cana 1:1—490; 2:1-622. 

ForBES, W. T. M. 1930. Insects of Porto Rico and Virgin Islands. Sci. Surv. Porto Rico 
and Virgin Islands. N.Y. Acad. Sci. 12:1-171. 

HABER, W. A. 1984. Pollination by deceit in a mass-flowering tropical tree, Plumeria 
rubra L. (Apocynaceae). Biotropica 16:269-275. 


214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


HopcEs, R. W. 1971. The moths of America north of Mexico. Fasc. 21. Sphingoidea. 
E. W. Classey, Ltd. and R. B. D. Publ. Inc., London. 158 pp. 

JANZEN, D. H. 1980. Two potential coral snake mimics in a tropical deciduous forest. 
Biotropica 12:77-78. 

1983. Pseudosphinx tetrio (Oruga Falso-Coral, Frangipani Sphinx). In Costa 
Rica Natural History. D. H. Janzen, ed. Univ. Chicago Press, pp. 764-765. 

MARTORELL, L. F. 1945. A survey of the forest insects of Puerto Rico. J. Agric. Univ. 
P.R. 39:1-354. 

1976. Annotated food plant catalog of the insects of Puerto Rico. Univ. P.R., 
Agric. Exp. Sta., Dept. Entomol. 303 pp. 

MEDINA-GAUD, S. & L. F. MARTORELL. 1974. The insects of Caja de Muertos Island, 
Puerto Rico. J. Agric. Univ. P.R. 58:244-272. 

Moss, REV. A. M. 1920. Sphingidae of Para, Brazil. Nov. Zool. 27:333-—424. 

PogEy, P. H. 1832 [1970]. Centurie de Lepidopteres de |'lle de Cuba. E. W. Classey, 
Ltd., London. 

WoLcoTT, G. W. 1948. The insects of Puerto Rico. J. Agric. Univ. P.R. 32:1-608. 








GENERAL NOTES 


Journal of the Lepidopterists’ Society 
39(3), 1985, 215-223 


NATURAL HISTORY NOTES ON ASTRAPTES AND URBANUS 
(HESPERIIDAE) IN COSTA RICA 


The close evolutionary affinity between Astraptes and Urbanus skippers (Hesperiidae) 
as members of the “Urbanus group” within the Pyrginae (Evans, 1952, Catalogue of the 
American Hesperiidae. Part II, British Museum of Natural History, London, 178 pp.) 
suggests similarities in the comparative biology of immature stages among representative 
species in both genera. For example, published larval food plant records for both genera 
include Leguminosae, and Astraptes has only been found feeding on members of this 
family (e.g., Howe, 1975, The butterflies of North America, Doubleday & Co., New 
York, 591 pp.; Kendall, 1976, Bull. Allyn Mus. No. 39, 9 pp.). While the majority of 
Urbanus species are legume-feeders as caterpillars, a handful of species are grass-feeders 
(Howe, op. cit.). Further field studies on the natural history of selected species in both 
genera may either confirm existing patterns of larval food plant patterns or augment 
them with records involving yet other families of dicotyledonous plants. It is evident 
that, within the Hesperiidae, tropical genera and species have undergone considerable 
evolutionary divergence in terms of larval food plants, with ten or more distinct food 
plant families known for some regions of the Neotropics (e.g., Kendall, op. cit.). In this 
note I summarize life cycle and larval food plant notes for A. fulgerator (Walch) in 
Costa Rica and make some brief comparisons with similar data on U. proteus (Linnaeus) 
from the same or similar localities within the country. Information on egg laying behavior 
is also presented. Two new larval food plant families for A. fulgerator are presented. 
The comparison of larval biology between these two genera was prompted by the striking 
differences in larval appearance between them, even when both are found on legume 
food plants and suggesting very different strategies of larval defense against visually 
hunting predators. 

Life cycle and related natural history notes on A. fulgerator and U. proteus were 
accumulated intermittently over more than ten years at the following Costa Rican lo- 
calities: “Bajo La Hondura” near Coronado (10°03'N, 84°00’W; 900 m elev.), San Jose 
Province (1972-1973); “Cuesta Angel” near Cariblanco (10°16’N, 84°10'W;; 1000 m elev.), 
Heredia Province (1973); “Finca La Tigra” near La Virgen (10°23’N, 84°07'W; 220 m 
elev.), Sarapiqui District in Heredia Province (1982); “Finca La Lola” near Siquirres 
(10°06'N, 83°30’W; 80 m elev.), Limon Province (1983); “San Rafael de Ojo de Agua” 
near Ojo de Agua (8°41'N, 83°28’W; 600 m elev.), Alajuela Province (1984); “Barranca 
Forest’ near Puntarenas (9°30'N, 84°35’W; 50 m elev.), Puntarenas Province (1984). 
These localities encompass a broad range of vegetational formations, from lowland to 
montane tropical wet forest, and including highly seasonal regions (Ojo de Agua and 
Barranca). All observations on egg laying behavior and collections of early stages were 
made in highly disturbed secondary habitats, including the borders of forest (La Tigra), 
forest remnants (Ojo de Agua and Barranca), and in a cacao plantation (La Lola). Im- 
mature stages were often reared through adulthood by confining eggs or caterpillars in 
tightly closed clear plastic bags containing fresh cuttings of the food plant. Food plant 
voucher specimens were collected for identification in all instances. Notes are also in- 
cluded for an undetermined species of Astraptes. 


Urbanus proteus Natural History 


Various authors have discussed the life cycle of this common skipper, and in this note 
I highlight only certain aspects of natural history relevant to a discussion of larval food 
plant exploitation and comparative behavior of immature stages between this species 
and A. fulgerator. Eggs (each about 1.8 mm dia., white to pale yellow with vertical 
grooves; spherical with flattened top) are deposited singly on ventral surface of mature 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Egg, first-instar, and second-instar caterpillars of Urbanus proteus in Costa 
Rica. Upper panel, from left to right: recently deposited egg, lateral view; egg in the 
process of hatching; egg shell after hatching. Bottom panel, left to right: first-instar about 
10 minutes after hatching; first-instar constructing shelter (note silk threads); second- 
instar feeding at the edge of a mature leaf of Mucuna sp. (Leguminosae). 


leaves of Mucuna spp. (Leguminosae), and only the top of the egg shell is eaten during 
the hatching process (Fig. 1). The first and second instar caterpillars construct a shelter 
in which to be concealed by folding over an irregularly shaped fragment of leaf along 
the leaf edge (Fig. 1). The caterpillar lines both surfaces of this structure with a loose 
network of silk and generally perches on the underside of the “roof” portion of the 
shelter (Fig. 2). Although initially a pale reddish brown, the caterpillar generally retains 
the same basic body color pattern throughout all instars: head capsule markedly bi-lobed 
vertically and shiny dark brown; neck “collar” red-brown above and red below (lateral- 
ventral flanges); body light green with “speckled” appearance and sparse pubescence of 
short, soft hairs (white); body sometimes appearing orange-green and with one pair of 
longitudinal orange lines running the length of the body; next-to-last abdominal segment 
with one pair of large orange blotches dorso-laterally; anal plate dark greenish brown 


VOLUME 39, NUMBER 3 217 


\ 
6 
_ 
a 
HI 
y 
7 
i 

ro 





Fic. 2. Urbanus proteus natural history. Upper panel, left to right: Mucuna leaves 
showing characteristic feeding pattern of young caterpillar; dorsal view of tent shelter 
of second-instar caterpillar on the dorsal surface of the leaf. Bottom panel, left to right: 
third-instar caterpillar perching on silk mat on the “roof” portion of tent shelter; fourth- 
instar tent shelter from dorsal aspect and with caterpillar hidden from view; fourth-instar 


caterpillar perched on “roof” portion of tent shelter shown in the previous photograph 
(match up areas of leaf damaged by larval feeding). 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 3. Dorsal, ventral, and lateral views of pupa of Urbanus proteus, respectively. 


but with thin border of orange. Later instars fashion tent shelters by tying together two 
lobes of the tri-lobed Mucuna leaf (Fig. 2); always solitary. Entire larval period lasts 
about 45 days, with transition from third to last (fifth) instar in about 20 days with an 
increase of 16 mm in body length (from 19 to 35 mm). The 21 mm long x 6 mm wide 
chestnut brown pupa is generously dusted with a bluish white pubescence (Fig. 3) and 
is formed with the tent shelter used by the final instar caterpillar. Eclosion takes place 
in about 16 days. In Costa Rica, caterpillars of U. proteus appear to be dusk or nocturnal 
feeders. It is very likely that this species occurs on a broad range of legumes in Costa 
Rica as noted elsewhere (e.g., Comstock & Vazquez, 1961, Ann. Inst. Biol. 31:349-448; 
Howe, op. cit.). 


Astraptes fulgerator Natural History 


Egg very similar to that of U. proteus except 2.0 mm diameter and placed on both 
meristem and mature food plant leaves. In a large patch of Erythrina sp. (Leguminosae) 
seedlings all less than 0.5 m tall in the cacao grove at La Lola, a fresh appearing female 
placed one egg each on a total of five seedlings within a two minute period one morning. 
In each case, the butterfly alighted on the underside of a large meristem leaf and quickly 
affixed an egg to it (12 March 1983). At La Tigra, one female distributed a total of eight 
eggs within a 7.0 m tall Erythrina tree over several minutes during a morning of inter- 
mittent drizzle and sunshine. All eggs were affixed singly to the undersides of mature 
leaves, one egg per leaf and widely scattered throughout the lower portion of the leafy 
canopy. During a drizzle spell, the butterfly flew off, only to return about ten minutes 
later to resume oviposition in this tree. Egg hatching involves devouring only the top 
portion of the shell, and the caterpillar (about 3 mm long with reddish brown body and 
large black head capsule) immediately crawls to the edge of the same leaf and constructs 
a tent shelter in a manner identical to that described earlier for U. proteus. The first 
and second instar larva rests on the underside of the “roof” leaf flap of the shelter; later 
instars build larger tents as also noted for U. proteus. Fifth instar larvae of A. fulgerator 
were found at Ojo de Agua (26 February 1984) feeding on mature leaves of Calea 
urticifolia (Willd.) (Compositae) and a single fifth instar of this species was found on 
Trigonia rugosa Benth. (Trigoniaceae) at Barranca a week later (2 March 1984). In both 
cases, larvae were concealed inside tent shelters similar to those found on Mucuna for 
an unidentified species of the same genus (see below) and for U. proteus. Both food plant 
shrubs possessed predominantly mature, well-worn, and heavily insect damaged leaves, 


VOLUME 39, NUMBER 3 219 


with little or no meristem leaves evident during this dry season period (January—April) 
at both localities. Mature caterpillars of A. fulgerator were present on both food plants 
at a time when meristem leaves were either entirely absent (T. rugosa) or very scarce 
(C. urticifolia). For C. urticifolia there was a mean length (x + S.D.) of 1.02 + 0.43 cm 
for meristem leaves (defined here as soft unfurling leaves within the length range of 0.2- 
4.0 cm) for a total of 18 leaves measured on three branches on 26 February 1984. On 
15 March 1984, x + S.D. was 2.54 + 1.05 cm for a total of 36 meristem leaves within 
the same size range on the same three branches. In captivity A. fulgerator fifth instars 
successfully completed development on the mature, worn leaves of their food plants. 
When the caterpillar and its woody vine-like food plant shrub were discovered along a 
foot path within the Barranca forest habitat (Fig. 4), a freshly eclosed adult A. fulgerator 
was netted about 10 m from the spot one hour later. 

The fifth instar caterpillar of A. fulgerator attains a body length of 48 mm, a maximal 
body width of 8 mm, and a head capsule width of 6 mm. The reddish brown, markedly 
bi-lobed head capsule is densely covered with short-to-long white hairs (Fig. 5). The 
neck “collar” is yellowish orange and the ground color of all body segments is a deep 
wine-red; each body segment with a thick transverse white band, and the entire body is 
blanketed with fine white hairs of varying lengths (Fig. 5). The anal plate is dull red as 
are all legs. During the daytime, the caterpillar remains well concealed in a shelter 
formed by folding over a portion of leaf and anchoring it with a few silk threads (Fig. 
5). The pupa is housed in a tent shelter formed by pulling together two or more adjacent 
leaves (Fig. 5). The pupa itself is very similar to that of U. proteus, but measuring 25 
mm long by 9 mm wide; the cuticle is generously covered with a dusting of bluish white 
pubescence (Fig. 5). Eclosion takes place in about 19 days. 


Unidentified Astraptes Natural History 


The fifth instar caterpillar stage of an unidentified Astraptes was found concealed in 
a tent shelter on a Mucuna vine at Bajo La Hondura (26 December 1972) (Fig. 6). The 
caterpillar was 835 mm long when discovered and grew to 50 mm in length by 18 January 
1973, at which time a massive number of larvae of an endoparasite emerged from it and 
formed a mass of cocoons on the cuticle (Fig. 6). The caterpillar is dark brown with 
conspicuous lateral blotches of pale green; the anal plate is dull red and the head capsule 
a glossy dark brown. The head capsule is covered with short reddish hairs. A second 
caterpillar found at this site was reared to the pupa stage. About two days prior to 
pupation, it became an active, orange colored prepupa, eventually pupating within its 
tent shelter. The 25 mm long by 8 mm thick pupa similar to those previously described. 
The parasitized caterpillar yielded a total of 100 Apanteles sp. wasps (Hymenoptera: 
Braconidae: Microgasterinae), a group known only to be endoparasites of Lepidoptera 
caterpillars (P. M. Marsh, pers. comm.). 

Astraptes fulgerator has been reported as feeding on various Leguminosae (e.g., Com- 
stock & Vazquez, op. cit.; Howe, op. cit.) and other species of the genus also on legumes 
(Kendall, op. cit.). The discovery in Costa Rica of this skipper exploiting both Compositae 
and Trigoniaceae as larval food plants is new to science. Kendall (op. cit.) lists some 11 
plant families as being reliable larval food plant records for Hesperiidae in Mexico, but 
that list does not include Compositae or Trigoniaceae. One plant family that does turn 
up in hesperiid larval food plant records in the American tropics and subtropics is Mal- 
pighiaceae, a group found along with Trigoniaceae in the order Polygalales of the subclass 
Rosidae (Cronquist, 1981, An integrated system of classification of flowering plants, Co- 
lumbia, New York, 1262 pp.). Furthermore, the Leguminosae, a common hesperiid larval 
food plant family and utilized by both Urbanus and Astraptes, is also within the Rosidae, 
but in a different order, the Fabales (Cronquist, op. cit.). Howe (op. cit.) summarizes the 
wide geographical distribution of A. fulgerator and the wide variability in the color 
pattern of the caterpillar stage. It would not be surprising to discover a polyphagous 
habit in such a species, and this note confirms this pattern for A. fulgerator in Costa 
Rica. While both Urbanus and Astraptes exploit legumes such as Mucuna vines in Costa 
Rica, the relatively aposematic appearance of Astraptes caterpillars compared with the 
subdued or cryptic-like colors of Urbanus caterpillars suggests a divergence in larval 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


| ce. Nor a i 
Fic. 4. Habitat and larval food plant of Astraptes fulgerator at Barranca. Top: 
disturbed primary-secondary forest where the larval food plant, Trigonia rugosa (Tri- 
goniaceae) was found. Bottom, left and right: habitat area where freshly-eclosed adult 


A. fulgerator netted; T. rugosa showing insect-damaged mature leaves (machete for 
scale). 





VOLUME 39, NUMBER 3 DADA | 





Fic. 5. Astraptes fulgerator natural history. Above panel, left to right: fifth-instar 
caterpillar, lateral view; tent shelter of fifth-instar caterpillar on Trigonia rugosa food 
plant. Bottom, left to right: tent shelter containing pupa; ventral aspect of pupa showing 
dusting of pubescence. 


defense against predators that possess color perception abilities. While the seeds of some 
Mucuna species possess toxic secondary compounds demonstrated to thwart predation 
by vertebrates (e.g., Janzen, 1969, Evolution 23:1-27), far less is known about the exis- 
tence of poisonous compounds within the leaves of these vines. Other herbivores routinely 
associated with some Mucuna species in Costa Rica, such as Morpho peleides Kollar 


222 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


‘ee 
Ese iaed “ha pi tae eee 


3 * 
<= om 
g ® % a 
a. = 
aa B 
f 


ee 





Fic. 6. Unidentified Astraptes species natural history. Above panel, left to right: tent 
shelter of fifth-instar caterpillar on Mucuna leaf and showing silken ties; close-up view 
of silken threads responsible for holding opposite sides of leaf together; tent shelter of 
an early instar at edge of leaf. Bottom panel, left to right: fifth-instar caterpillar, lateral 
view; mass of Apanteles cocoons on surface cuticle of fifth-instar caterpillar. 


(Morphidae), have brightly colored early instar caterpillar stages and adults with irides- 
cent blue wings (e.g., Young & Muyshondt, 1978, Carib. J. Sci. 18:1—49). In some lepi- 
dopterans associated as herbivores with plants possessing known toxic properties, some 
species are able to sequester the toxic compounds, while others, feeding on the same 


VOLUME 39, NUMBER 3 293 


plants, are not able to do so (e.g., Rosenthal & Janzen, eds., 1979, Herbivores: Their 
interaction with secondary plant metabolites, Academic Press, New York and London, 
717 pp.). Thus, the possibility exists that Astraptes, with brightly colored caterpillars and 
adults with iridescent blue wings, have evolved the ability to sequester presumed toxins 
in the leaves of legume food plants such as Mucuna, while Urbanus has not evolved such 
a trait. Further support for a presumed toxicity associated with Astraptes is afforded by 
the Calea urticifolia larval food plant record reported here: this plant is known to be 
extremely toxic owing to high concentrations of sesquiterpenes (L. Poveda, pers. comm. ). 
To the best of my knowledge, Urbanus does not feed on Compositae or Trigoniaceae. 

The discovery of mature caterpillars of Astraptes at the height of the tropical dry 
season at two localities indicates that these insects are able to exploit mature leaves at 
such times of the year when meristems are lacking or scarce. The cohort of adult skippers 
eclosing in the latter half of the dry season might “anticipate” a soon-to-be available 
supply of fresh meristems on which to place their eggs in the sense of Phoebis exhibiting 
such behavior on its Cassia food plant at this time (Young, in press). My data suggests 
that Astraptes will deposit eggs on both mature leaves and meristem leaves. To what 
extent, if any, does A. fulgerator exhibit a facultative seasonal switch in food plant 
families in Costa Rica awaits further field study. To what extent does Mucuna and other 
legume food plants, fully leafed-out in the rainy season, become inaccessible as larval 
food sources during the tropical dry season and inducing a switch to alternate food plant 
groups, remains to be studied. Food plant quality, and temporal changes in it, is a major 
determinant of food plant choice in some herbivorous insects (e.g., Marian & Pandian, 
1980, Entomon 5:257-264; Rausher, 1981, Ecol. Monogr. 51:1-20; Hill, 1982, Zool. Jahrb. 
Abt. Syst. Okol. Geogr. Tiere 109:24-32; Lawson et al., 1982, Entomol. Exp. Appl. 32: 
242-248; Messina, 1982, Oecologia 55:342-354; Miles et al., 1982, Aust. J. Zool. 30:347- 
355; Wint, 1983, J. Anim. Ecol. 52:438-—450). Some skippers exhibit seasonal shifts in the 
use of their larval food plants (e.g., Nakasuji, 1982, Appl. Entomol. Zool. 17:146-148). A 
polyphagous insect such as A. fulgerator in Costa Rica may possess the complement of 
mixed-function oxidases within the guts of caterpillars to permit the expression of a 
generalist feeding behavior involving different food plant families exhibiting both qual- 
itative and quantitative differences in the profiles of toxic compounds functioning to 
deter herbivorous attack (e.g., Ahmad, 1988, Ecology 64:235-243). Finally, within a 
relatively small region of the American tropics, a polyphagous species such as A. fulger- 
ator may have undergone an evolutionary divergence in the use of different larval food 
plant families. Sometimes such geographical divergence reaches the point at which cat- 
erpillars from different populations cannot survive on food plants other than those found 
in their own habitats (e.g., Kaufmann, 1983, Proc. Entomol. Soc. Wash. 85:321-326). I 
was unable to detect any noticeable difference in coloration between the caterpillars of 
A. fulgerator from Ojo de Agua and Barranca, and the sample was very small. Astraptes 
caterpillars may use tent shelters on their food plants as a means of hiding from parasit- 
oids such as Apanteles and Tachinidae, but such behavior does not always ensure survival 
as noted in this paper. 

This work was partially supported by a grant from the Friends of the Museum (of the 
Milwaukee Public Museum), the National Science Foundation, and the American Cocoa 
Research Institute. Luis Diego Gomez and Luis Poveda (Museo Nacional de Costa Rica) 
provided determinations of larval food plants and additional comments on the biology 
of these plants. Stephen R. Steinhauser graciously brought to my attention the important 
Kendall paper. P. M. Marsh (U.S. National Museum of Natural History) provided the 
determination of the parasitic wasp. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 532833. 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Journal of the Lepidopterists’ Society 
39(3), 1985, 224-225 


INDEPENDENT EVOLUTION OF “FALSE HEAD” 
BEHAVIOR IN RIODINIDAE 


Although “false head” wing patterns and behaviors, particularly hindwing movements 
along the sagittal plane, are well-known among Lycaenidae (Robbins, 1980, J. Lepid. 
Soc. 34:194-208; 1981, Am. Nat. 118:770-775), it is less well-documented that some 
Riodinidae have similar wing patterns. It is reported here for the first time that some 
have also independently evolved hindwing movements similar to those of lycaenids. This 
situation is intrinsically interesting as an example of convergent evolution and is addi- 
tionally significant for its phylogenetic implications. 

Probably the best-developed riodinid “false heads’ occur in Helicopis Fabricius and 
some species of Sarota Westwood. These butterflies have multiple white-tipped tails and 
metallic markings at the anal angle of the ventral hindwing. They land with their wings 
folded over their backs, unlike many riodinids, and are often mistaken for Lycaenidae. 
Maj. Harold Harlan (Ohio State Univ., pers. comm.) first noted a specimen of Sarota (a 
species with hindwing tails) (Canal Zone, Panama) moving its hindwings while landed. 
In San Carlos de Rio Negro (Amazonas, Venezuela), I observed 11 specimens of Helicopis 
cupido erotica Seitz and two specimens of Anteros formosus Cramer (a species that lacks 
hindwing tails) moving their hindwings. Curtis J. Callaghan (Petropolis, Brasil, pers. 
comm.) has also observed this behavior in Helicopis, Sarota, and Anteros Hiibner (and 
noted in addition that the long tails of Helicopis are easily moved by breezes). Hindwing 
movements were previously reported only among Lycaenidae. 

Hindwing movements in riodinids, as in lycaenids, may occur sporadically. The lit- 
erature on lycaenid behavior includes cases where one observer noted hindwing move- 
ments, whereas, another did not in the same species under similar circumstances (Rob- 
bins, 1980, op. cit.). The same appears to be the case in riodinids. Whereas, I did not 
observe specimens of Panamanian Sarota moving their hindwings, Harlan did. Whereas, 
I noted individuals of Anteros formosus moving their hindwings in southern Venezuela, 
Callaghan did not when he came upon a swarm of this species in Brasil’s Mato Grosso. 
These examples point out the difficulties of interpreting negative evidence with regard 
to sporadically occurring behaviors. 

Although “false head’’ wing patterns and behaviors of lycaenids and riodinids are 
superficially similar, they differ in detail. Lycaenid anal lobes are everted outwards while 
tails project inwards and cross (see fig. 12 in Longstaff, 1912, Butterfly-hunting in many 
lands, Longmans, Green, and Co.). In contrast, both tails and anal lobes of Helicopis 
flare outwards. Hindwing movements also differ. Lycaenids move both hindwings si- 
multaneously; as one hindwing moves forward, the other moves backwards, and vice 
versa. In contrast, Helicopis and Anteros may move one or both hindwings. If both are 
moved, they need not be in opposite directions. In addition, their movements are “jerky” 
and of short duration in contrast to lycaenid movements, but I saw too few specimens 
to quantify this difference. These morphological and behavioral differences support taxo- 
nomic evidence that the “false heads” of lycaenids and riodinids are independently 
evolved. 

The distribution of lycaenid “false heads” are phylogenetically significant. Eliot (19783, 
Bull. Brit. Mus. (Nat. Hist.) Entomol. 28:6) suggested that Theclinae, Polyommatinae, 
and Lycaeninae form a monophyletic clade. Among the Lycaenidae, “false heads” are 
known only in these subfamilies and are the only characters of which I am aware that 
are unique to them. This being the case, it is desirable to have better documentation of 
the distribution of hindwing movements within the Lycaenidae, particularly among the 
many Old World tribes. 

Riodinid “false heads” are also phylogenetically significant. Stichel (1930-1931, Lep- 
idopterorum catalogus, pars 40) and Clench (1955, Annals Carnegie Museum 33:261- 
274) placed Helicopis in its own tribe. Harvey (Univ. Texas, Austin, dissertation in prep.), 


VOLUME 39, NUMBER 3 225 


on the other hand, proposes that Helicopis, Sarota, Anteros, and Ourocnemis Baker form 
a closely related group of genera. The behavioral evidence reported here supports Har- 
vey s classification. Further, we can predict that specimens of Ourocnemis will be found 
to also move their hindwings. 

I thank C. J. Callaghan, J. H. Harlan and D. J. Harvey for permission to report their 
findings and for critically reading the manuscript. I acknowledge a Scholarly Studies 
Grant for supporting and the Fundacion para el Desarrollo de las Ciencias Fisicas, Ma- 
tematicas y Naturales for sponsoring the field trip on which these observations were 
made. 


ROBERT K. ROBBINS, Department of Entomology, MRC NHB 127, National Museum 
of Natural History, Smithsonian Institution, Washington, D.C. 20560. 


Journal of the Lepidopterists’ Society 
89(3), 1985, 225-228 


INTERACTIONS OF PARASITOIDS WITH AN OPSIPHANES 
(BRASSOLIDAE) CATERPILLAR IN COSTA RICA 


Both hymenopteran and dipteran parasitoids are known to kill the caterpillars of 
Opsiphanes species (Brassolidae) in Central America (Harrison, 1963, Ann. Entomol. Soc. 
Amer. 56:87-94; Young & Muyshondt, 1975, Stud. Neotrop. Fauna 10:19-56). In these 
studies, wild-caught caterpillars on their monocot food plants (Musaceae and Palmae) 
are checked individually for emergence of parasitoids, with little or no direct observations 
on the ways in which these organisms interact with their host. In this note I describe 
some behavioral observations on both adult Tachinidae (Diptera) and Chalcidae (Hy- 
menoptera) attempting to parasitize a single Opsiphanes caterpillar at the same time. 
Field observations on the interaction of hymenopteran and dipteran parasitoids on the 
same host are almost entirely absent in the entomological literature. Given the well- 
documented and studied roles of individual parasitoid species in the regulation of plant- 
associated insect populations in both the temperate and tropical zones (e.g., Jumalon, 
1964, J. Lepid. Soc. 18:101-104; Herrebout, 1966, Z. Angw. Entomol. 58:340-355; Etch- 
egaray & Nishida, 1975a, Proc. Hawaiian Entomol. Soc. 22:33-39; 1975b, Proc. Hawaiian 
Entomol. Soc. 22:41-49; Link, 1977, Dusenia 10:201-204; Zaucki, 1981, Aust. Entomol. 
Mag. 8:3-8; Olaifa & Akingbohungbe, Insect Sci. Appl. 3:73-77; Roth et al., 1982; En- 
viron. Entomol. 11:273-277; Stamp, 1982, Environ. Entomol. 11:100-104; Courtney & 
Duggan, 1988, Ecol. Entomol. 8:271-278; Elzen et al., 1983, Environ. Entomol. 12:1872- 
1876; Grant & Shepard, 1983, Environ. Entomol. 12:1673-1677; Messina, 1983, Environ. 
Entomol. 12:807—809; Oatman et al., 1983, J. Econ. Entomol. 76:52-53; Thompson et al., 
1983, Environ. Entomol. 12:1312-1314; Maier, 1984, Can. Entomol. 116:443-449; Mar- 
ston et al., 1984, Ann. Entomol. Soc. Amer. 77:21-28), the field study of the ways in 
which different parasitoids “interact” at the same host may clarify certain aspects of how 
these organisms regulate populations of phytophagous species (e.g., Van Driesche, 1983, 
Environ. Entomol. 12:1611-1622). 

A fifth instar caterpillar of Opsiphanes sp. was found partly concealed within a silken 
sleeve on a single pinna of a coconut palm (Cocos nucifera “dwarf” variety) on 7 March 
1984 at “Finca Experimental La Lola,” about 15 km east of Siquirres (10°06’N, 83°30'W), 
Limon Province, Costa Rica. From 7 to 16 March, I made daily observations at various 
hours on the presence of tachinids and hymenopterans with this caterpillar. These ob- 
servations were initiated when, at 1745 h on 7 March, I observed a tachinid (described 
as ‘red eyes with gray and white striped body and wings held at about 45° angle to the 
body”) “buzzing” around the caterpillar as the latter crawled towards the silken sleeve 
from an apparent feeding site elsewhere on the tree (about 2.0 m tall). On 16 March the 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


caterpillar was collected and kept alive for rearing to adulthood, placing it in a clear 
plastic bag with a fresh cutting of the coconut palm leaf pinnae. 

Between 7 and 12 March, the Opsiphanes caterpillar was found re-entering its “nest” 
on the same pinna, usually between 1600 and 1800 h; morning observations (0800-1100 
h) on the same days indicated no activity; the caterpillar could clearly be seen resting 
motionless inside the nest. The nest was located on the distal-most third of the pinna, 
and the caterpillar either crawled directly into it, head-first (Fig. 1) or sometimes backed 
into it. 

A single adult tachinid was observed attempting to land on the caterpillar on two 
dates: 7 March (1745 h) and 12 March (1500 h). Based upon the general appearance of 
the fly, it appeared to be the same species on both dates and it was collected on 12 
March, following observations, and kept for a voucher determination. On 7 March, the 
fly made several attempts to alight directly on the caterpillar’s body, usually in the 
proximal-most third region. Every time the fly made such an attempt (N = 18 instances) 
to land, the caterpillar jerked its head violently in broad swings through an imaginary 
circular axis, effectively chasing the fly away for a few moments. Between attempts, the 
tachinid often alighted on an adjacent pinna. For a total of 11 times, the fly intermittently 
perched alongside the caterpillar as it crawled towards the nest (Fig. 1), periodically 
trying to alight on the host again but always failing to do so. Once inside the nest, the 
tachinid walked over the silken surface (Fig. 1) of it, before flying off (by 1755 h). On 
12 March at 1500 h, a tachinid was found walking on top of the silken nest again, when 
the caterpillar was motionless inside. At one point, the fly sat motionless immediately 
above the thoracic area of the caterpillar’s body but on the outside of the thin silken 
layer between it and the host insect. The caterpillar was clearly visible through the fine 
lacework of silken mesh (Fig. 1). Periodically, the fly moved a bit and curled its abdomen 
under itself, as if attempting to oviposit either on the silk or through it and onto the body 
of the host. At 1700 h, I gently placed a dry glass vial over the tachinid as it perched 
above the caterpillar, and it did not move at all. The fly was collected in this manner a 
few minutes later. 

On 11 March at 1545 h, I noticed a small black wasp securely fastened to the dorsal 
area of the thorax of the caterpillar. In spite of violent, head-thrashing movements 
identical to those exhibited in an apparent response to the presence of the tachinid, the 
caterpillar was unsuccessful in dislodging this wasp. At the time the caterpillar was about 
three cm from the entrance of the silken nest. It moved into the nest with the wasp still 
firmly attached. Once inside the nest, the wasp began crawling over the thoracic area of 
the Opsiphanes caterpillar. A few moments later, the big caterpillar backed out of the 
nest with the wasp still attached. At this time, in the good daylight, I noticed a few 
blackened spots on the caterpillar’s thoracic area, easily spotted against the light green 
background color of the insect. The caterpillar, however, was not successful in removing 
the wasp. I collected the wasp a few moments later (using a glass vial). 

The Opsiphanes caterpillar was collected at 1800 h on 12 March for rearing. Two 
days later, three tachinid pupae were found in the bottom of the rearing bag and cat- 
erpillar was dead. About a week later the tachinids eclosed and they matched the general 
appearance of the adult observed interacting with the caterpillar in the wild. No parasitic 
hymenopterans emerged. 

The four tachinids were examined a few weeks later by Dr. Norman E. Woodley, 
Research Entomologist, Systematic Entomology Laboratory of the U.S. Department of 
Agriculture (Washington, D.C.), who told me that they were from a very poorly known 
group, making generic determination exceedingly difficult. The wasp was determined to 
be Brachymeria sp. (Hymenoptera: Chalcidae) by Dr. E. E. Grissell of the U.S. National 
Museum of Natural History. Species determination of the Opsiphanes was not confirmed 
since the adult was not reared. 

These qualitative observations suggest that Opsiphanes caterpillars exhibit two types 
of defensive behavior against parasitoids and do so in the following order: (1) violent 
head-thrashings chase away tachinids some times, followed by (2) rapid movement into 
a silken nest. If a caterpillar is approached by a parasitoid some distance away from the 
nest, the second line of defense cannot be utilized. Given the fact that the caterpillar 


VOLUME 39, NUMBER 3 yagegt 





Fic. 1. Left: Opsiphanes caterpillar entering silken nest on leaf pinna of the food 
plant, Cocos nucifera (Palmae) at Finca Experimental La Lola, Siquirres, Limon Prov- 
ince, Costa Rica (12 March 1984 at 1600 hr); right: adult tachinid perched on the silken 
“roof” of the caterpillar nest and directly over the thoracic region of the host’s body (12 
March 1984 at 1550 h). 


died from tachinid attack, it is clear that the defensive behavior is not always successful. 
The close association of both kinds of parasitoids with the caterpillar at its nest site further 
suggests that chemical attractants associated with the nest itself might be involved in the 
host-searching behavior of the parasitoids. Volatiles are known to play important roles 
as chemical signals in attracting parasitoids to their hosts (e.g., Weseloh, 1980, Ann. 
Entomol. Soc. Amer. 73:593-601; Elzen et al., op. cit.; Kamm & Buttery, 1983, Entomol. 
Exp. Appl. 33:129-134; Thompson et al., op. cit.), although physical (structural) stimu- 
lants may also be involved in some instances (e.g., Cole, 1959, J. Lepid. Soc. 13:1-10; 
Tautz & Markl, 1978, Behav. Ecol. Sociobiol. 4:101-110). The possible role of lepidop- 
teran silken nests as deterrents to parasitoids has not been studied in great detail, although 
Stamp (op. cit.) observed parasitoids attacking the gregarious caterpillars of the Baltimore 
_checkerspot on the outside of their nests. The nesting habit of Opsiphanes has been 
known for some time (Jones, 1882, Proc. Liter. and Philosoph. Soc. Liverpool 36:327- 
377), but the functional role of this habit remains largely unstudied. 

Young and Muyshondt (op. cit.) noted that O. tamarindi Fruhstorfer is parasitized by 
two different species of Tachinidae in El Salvador and Costa Rica, and it was assumed 
in that paper that adult parasitoids deposited their eggs on the food plants rather than 
on the host caterpillar. But one caterpillar of O. tamarindi in El Salvador was found 
with two eggs attached to the integument near the distal end of the trunk, and these 
might have been Tachinidae. Young and Muyshondt (op. cit.) found considerable para- 
sitism of O. tamarindi by the braconid Meteorus sp. in E] Salvador and by the chalcid 
Spilochalcis nigrifrons (Cam.) in Costa Rica. Ten of 11 pupae reared from wild-caught 
fourth and fifth instar caterpillars from Puntarenas, Puntarenas Province, Costa Rica 
eclosed as S. nigrifrons instead of adult O. tamarindi in that study. Tortricid and geo- 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


metrid caterpillars associated with avocado foliage in southern California frequently 
exhibit “mixed” parasitism from both braconids (Apanteles) and chalcids (Meteorus) 
(Oatman et al., 1983, op. cit.). Young and Muyshondt (op. cit.) suggest that tachinids 
associated with Opsiphanes populations in Central America are most likely generalists 
on a broad range of lepidopterous hosts. Whether or not hymenopterous and dipterous 
parasitoids converging ecologically on the same individual host caterpillar actually en- 
gage in competition for the host remains to be studied quantitatively. And the Opsi- 
phanes x Musaceae and Palmae interaction in Central America might be a good model 
system for such studies, given (1) the large body-size of the host caterpillars, (2) the 
exploitation of the caterpillars by braconids, chalcids, and tachinids (including the same 
individual host), (3) the relatively restricted monocot food plant association of the cat- 
erpillars, and (4) the apparent economic importance of some Opsiphanes (e.g., Harrison, 
op. cit.). 

Other brassolids associated with Palmae in Central American forests may not be ex- 
periencing the same forms of selection pressure from parasitoids as Opsiphanes. Com- 
munally nesting aggregations of the caterpillars of Brassolis isthmia (Bates), which con- 
struct leaf and silken nests from adjacent palm leaf pinnae and which exhibit a strongly 
crepuscular feeding activity outside the nest, do not, for example, experience attacks 
from parasitoids such as Tachinidae and Chalcidae (A. M. Young, unpubl. obsery. and 
field data). 

This research is a by-product of a grant from The American Cocoa Research Institute. 
I thank Drs. E. E. Grissell and Norman E. Woodley (U.S. National Museum of Natural 
History and the Systematic Entomology Laboratory of the U.S. Department of Agricul- 
ture, respectively) for making determinations of the parasitoids mentioned in this paper. 
Parasitoid specimens are deposited in the national collections. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 532838. 


Journal of the Lepidopterists’ Society 
89(3), 1985, 228-229 


ON A PREVIOUS REPORT OF DIURNAL ROOSTING OF THE 
PIPEVINE SWALLOWTAIL, BATTUS PHILENOR (L.) 


Gillaspy and Lara (1984, J. Lepid. Soc. 38:142-143) recently recounted their obser- 
vations concerning a short-lived aggregation of Battus philenor (L.) in apparent response 
to an approaching rainstorm. They reported six butterflies flew to a branch of a mesquite, 
Prosopis glandulosa Torr., on 12 June 1981 near Laredo, Texas. Gillaspy and Lara (op. 
cit.) wondered if these butterflies would later return for a nocturnal roost, but they were 
unable to provide further observations. They also suggest that further, admittedly for- 
tuitous, observations would be required to understand fully such temporary behavior. 

Interestingly, observations by H. B. Parks (1935, Bull. Brooklyn Entomol. Soc. 30:196) 
on a similar occurrence provided some answers to questions raised by Gillaspy and Lara 
(op. cit.). On 7 June 1935, near Santa Rita in southern Brooks Co. (approximately 150 
km southeast of Laredo), he observed 40 B. philenor (prior to commencement of rain) 
fly toward and hang underneath the limbs of a huisache, Acacia smallii Isely, with, “The 
thick leaves and branches thus giving complete protection.” Of particular significance to 
one query presented by Gillaspy and Lara (op. cit.) is the report of a ““Ranchman [who] 
stated that these butterflies came to this one tree . . . also to roost during the night.” 

Field workers in the southern Texas area should be aware of the need for observations 
on these nocturnal roosts, if they exist. Obviously, each butterfly rests at night at some 


VOLUME 39, NUMBER 3 229 


location. The significant question is whether these butterflies rest individually or clumped 
in aggregations. 


RAYMOND W. NECK, Texas Parks and Wildlife Department, 4200 Smith School Road, 
Austin, Texas 78744. 


Journal of the Lepidopterists’ Society 
89(3), 1985, 229-235 


NATURAL HISTORY NOTES FOR SOME HAMADRYAS BUTTERFLIES 
(NYMPHALIDAE: NYMPHALINAE; AEGERONINI) IN NORTHWESTERN 
COSTA RICA DURING THE TROPICAL DRY SEASON 


The relatively small cluster of species belonging to the nymphaline genus Hamadryas 
are well known as ‘calicoes” or “crackers” in the adult stage throughout much of Central 
America, Mexico, and South America. The medium-sized gray-and-white speckled but- 
terflies are pugnacious, fast-flying insects that commonly perch on the trunks of trees 
during the daytime, head downwards, and with the wings held pressed down in the open 
position. Their fast aerial antics coupled with loud clicking noises evident in both sexes, 
and often involving attack approaches to “intruders” into their areas, have made them 
the subject of behavioral studies (e.g., Ross, 1963, J. Res. Lepid. 2:241-246). As a group, 
the caterpillars feed on Euphorbiaceae, particularly vines and shrubby plants of the genus 
Dalechampia (e.g., Young, 1974, Z. Angew. Entomol. 76:380-898; Muyshondt & Muy- 
shondt, Jr., 1975a, J. New York Entomol. Soc. 83:157-169; 1975b, J. New York Entomol. 
Soc. 83:170-180; 1975c, J. New York Entomol. Soc. 83:181-191; Jenkins, 1983, Bull. Allyn 
Mus. 81:1-146) and may function as significant selective agents in the evolution of 
herbivore resistance in these plants (Armbruster, 1982, Amer. J. Bot. 69:1429-1440). 
Adult butterflies are typically associated with open pastures and borders of dense vege- 
tation (Ross, 1964, J. Res. Lepid. 18:11-26; 1967, J. Res. Lepid. 18:11-26; 1967, J. Res. 
Lepid. 15:109-128; Monroe et al., 1967, J. Lepid. Soc. 21:185-197). Because some species 
of Hamadryas occur in the seasonal tropical dry forest zones of Central America, they 
offer the chance to study the impact of tropical seasonality upon their natural history. 
In this note we report such preliminary field studies from the lowland tropical dry forest 
zone of northwestern Costa Rica as performed during the dry season. Herein, we describe 
some hitherto unreported features of adult behavior, including nocturnal perching rel- 
ative to daytime perching, and evidence that, although females are mated during this 
period, they appear to be in a state of reproductive diapause. 

From 2-4 March 1984 we studied and collected adult Hamadryas from these two 
localities: (a) about 1.5 km south of Liberia (10°40’N, 85°40’W), Guanacaste Province 
and along the Pan American Highway, and (b) “Barranca Site” (Orians, 1969, Ecology 
50:783-801) about 6 km from Miramar (10°06’N, 84°44'W), Puntarenas Province. Both 
localities fall within the region of lowland tropical dry forest and experience a completely 
dry (no rainfall) season generally between December and May each year. Within a wide 
rectangular roadside area (approx. 50 m x 100 m) at the Liberia site, we studied the 
abundance and habits of adult Hamadryas on several mature forest canopy trees, mostly 
Guazuma ulmifolia Lam (Sterculiaceae). Approximately 75% of this area was covered 
by a dense patch of disturbed forest, consisting chiefly of a Guazuma “canopy” and 
fairly evergreen understory consisting of various Leguminosae, Flacourtiaceae, and other 
small trees and shrubs in varying degrees of “‘leafing out” at the time. We examined the 
distribution of adult Hamadryas perching on tree trunks within the forest patch and 
along its borders at various times of the day and night. As it was quickly apparent that 
the butterflies were most numerous along the strip of shade trees between the forest 
patch and the highway (Fig. 1), we concentrated our observations to that area which 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


contained eight mature G. ulmifolia, one Enterolobium cyclocarpum Donn. Smith. (Le- 
guminosae) and two palms (unidentified). We observed the occurrence of Hamadryas 
adults on the trunks of these and other trees during the daytime and also on two evenings; 
daytime observations were made intermittently from about 0800 and 1800 h on the two 
days. Evening or nocturnal observations were limited to 2000 to 2200 h. We searched 
tree trunks and foliage for resting butterflies at night, using head lamps and flashlights. 
On 3 March, we studied the movement of butterflies from tree trunks to other perching 
sites at 1700 to 1800 h. 

Potential adult food sources were noted as well as the availability of Dalechampia 
within the immediate vicinity (the forest patch) to determine the presence of immature 
stages. On the first night of study (2 March), we collected a series of adults for species 
vouchers, and during the day we made every attempt to recognize different “morpho- 
species” of adults and later matched vouchers with these field observations. We measured 
air temperature patterns in both shaded and exposed areas during the late morning, as 
well as wind strengths. To do the latter, we determined the degree to which the bag of 
a standard insect aerial net would be inflated (held horizontally) at the edge of the forest 
acting as a wind break and also in the adjacent pasture. We did this to determine a 
possible relationship between adult perching sites and protection from the strong wind 
gusts characteristic of the Guanacaste dry season. 

Three piles of rotting bananas were placed on the ground at the “Barranca Site” 
locality, a patch of semi-deciduous forest; these baits were placed along the foot path 
and the upper or southern end of this forest patch to attract adult Hamadryas. These 
three baits were scattered at 50-100 m intervals along the train, and the first one was 
placed at the base of a large Samanea saman (Leguminosae) tree, which is located about 
10 m from a small grove of G. ulmifolia trees. The baits were distributed at about noon 
on 2 March and reexamined for butterflies between 1400-1600 h on 4 March. Our 
purpose was to determine the Hamadryas species active here to compare with Liberia 
specimens, and to collect females for evidence of mating and reproductive condition. 

Adult female butterflies were stored in glassine envelopes and placed in a freezer upon 
return to Milwaukee a few days after the field work. The butterflies were then thawed 
and examined with a dissecting microscope to determine (a) the presence of spermato- 
phores and mating plugs, and (b) the degree of development of ovary tissues. The number 
of ova, immature and mature, were counted. Species determinations were made follow- 
ing the keys of Jenkins (op. cit.). 

Notes were taken on the presence and activity of other Papilionoidea at both localities 
to compare with Hamadryas. 

At about 2100 h on 2 March we collected five Hamadryas guatemalena guatemalena 
(Bates), and two H. glauconome glauconome (Bates) and one H. feronia farinulenta 
from the foliage of two adjacent trees along the front edge of the forest patch facing the 
highway. Other butterflies collected perching in the same foliage included Callicore 
pitheas (Latreille) and Opsiphanes cassina fabricii (Boisduval) (Brassolidae). Unlike day- 
time perching on tree trunks, nocturnally perching Hamadryas are positioned on the 
undersides of leaves and with their wings tightly folded. The same is true for other 
butterflies found perching on foliage at night (see also Young, 1979, J. Lepid. Soc. 33: 
58-60). In some instances, Hamadryas adults at night could be collected using fingers 
rather than a net. 

On the following afternoon, we documented the movement of Hamadryas from tree 
trunk perching sites to nearby foliage for nocturnal roosting. Between 1741 and 1756 h, 
we observed a total of 13 butterflies (three species: H. guatemalena guatemalena, H. 
glauconome glauconome and H. feronia farinulenta) fly into a single evergreen bush 
within 3-5 m of nearby G. ulmifolia trees used as daytime perching sites. At approxi- 
mately 1 to 2 minute intervals, 1-2 butterflies fluttered into the foliage from the sur- 
rounding area, and by 1758 h, all 13 individuals were perched on the undersides of leaves 
with wings closed, and all within 15 cm to 1.0 m of one another in the bush. Throughout 
the study period, the weather was hot, sunny and dry. During the dusk settling process, 
arrivals of some individuals resulted in others being temporarily disturbed, flying off and 


VOLUME 39, NUMBER 3 Zar 





Fic. 1. Clockwise, from upper left photograph: the western edge of the forest habitat 
at the Liberia locality—-Hamadryas was commonly found perching on the trunks of the 
trees to the right during the day and on the foliage to the left (4 March 1984); H. 
guatemalena resting on the trunk of a tree at the Liberia locality; Guazuma ulmifolia 
fruits on the ground; G. ulmifolia trees in the open pasture immediately south of the 
forest patch at the Liberia locality (also 4 March 1984). 


eventually returning. A check later that evening (about 2030 h) revealed no further 
additions. Butterflies did not perch on foliage during the daytime. 

A search for Hamadryas butterflies on tree trunks throughout the study site at Liberia 
(4 March at about 0900 h) revealed none associated with trees within the forest, nor any 
along trees bordering the eastern, northern, and southern boundaries of this patch; all 
butterflies were found on a few trees along the western border of the patch. Of eight G. 
ulmifolia trees in this area, only three had one or more Hamadryas perching on them 
during the day. There was almost an equal number of both sexes, judging from field 
observations and collections made, but one species, H. guatemalena, was more commonly 
encountered than others at both localities (Table 1). Most of these butterflies appeared 
to be “fresh” in terms of the wing condition, and all but one female was mated (Table 
1). Interestingly, only one species, H. guatemalena, was found at both localities, with an 
additional four species being distinct between them (Table 1). 

We could not detect any species-specific differences in daytime or nocturnal perching 
behavior at Liberia. It appeared that all species were responding to prevailing environ- 
mental conditions in the same manner. Mid-morning measurements (1000 to 1030 h on 
3 March) of air temperature in the shade at a G. ulmifolia tree frequently used for 
perching by Hamadryas and another individual of the same tree a few meters away and 
in direct sunlight revealed no differences (31.0°C at both trees at 1000 h and 30.6°C and 


232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Sex ratios, relative wing condition, and reproductive states for small samples 
of adult Hamadryas species at two localities in northwestern Costa Rica during the 
tropical dry season.* 








No. of adults 
Reproductive 


Date(s) Species Male Female Total Wing condition** state 





“Liberia, Guanacaste” 


2-4 March UH. guatemalena 4 ] 5 2mF, 2ml, 1fF mated 
1984 guatemalena (Bates) 
H. glauconome 2 0 2 2mF — 


glauconome (Bates) 
H. feronia farinulenta I 0 1 lmF _ 


(Fruhst. ) 
“Barranca Site, Puntarenas’ 
4 March H. guatemalena 0 2 2 2fF mated; 
1984 guatemalena (Bates) mated 
H. februa ferentina 0 1 I 1fF unmated 
(Godart) 
H. iphthime ipthime 0 il 1 1fF mated 
(Bates) 


Total species (both localities): 5 
Total individuals & overall sex ratio: 7 males + 5 females = 12 

* Counts were made of adult butterflies found perched on trunks, branches, and leaves of trees at the study sites. 
Other species present at the “Liberia” locality trees were: Callicore pitheas (Latreille), Eunica malvina (Bates), Siderone 
marthesia (Cramer), and Opsiphanes cassina fabricii (Boisduval). Reproductive states were examined for female but- 
terflies only. 


** T owercase letters refer to sex (m = male, f = female) while uppercase letters designate wing condition: F = fresh, 
I = intermediate-worn. 


30.8°C for exposed and shaded trees, respectively, at 1030 h). We did discover that wind 
gusts quickly inflated the bag of the aerial insect net along the open pasture area im- 
mediately adjacent to the forest patch, but the bag remained almost completely deflated 
when positioned along the western edge of the forest at the point where butterflies were 
perching on tree trunks (tests performed 0930-1000 h on 3 March). This observation 
suggests a wind-sheltering factor in the choice of tree trunks by Hamadryas. 

Counts of freshly fallen, sweet-smelling, G. ulmifolia fruits in four different one-by- 
one meter plots gave the following results: 70 and 73 fruits for two adjacent trees along 
the shaded western edge of the forest patch and 25 and 15 fruits each for two trees in 
the open pasture south of the forest patch. Since some butterflies in this region of Costa 
Rica feed on fallen G. ulmifolia fruits (e.g., Young, 1975, Rev. Biol. Trop. 23:101-128), 
we attempted to observe possible feeding by Hamadryas on these fruits at the Liberia 
locality, but this behavior was not observed. We searched for evidence of adult Hamadry- 
as feeding on these fruits, fresh cattle dung, and sap flows at the Liberia locality, but 
none was seen in spite of checking at various hours of the day. Most of these observations 
took place after 0830 h and we might have, therefore, missed an early morning feeding 
period. Of four Hamadryas observed near one of the banana baits at the Barranca Site, 
one of these was found feeding on the bait (1400 h, 4 March) along with several Caligo 
memnon Felder; three of the Hamadryas were perched on the S. saman tree immedi- 
ately behind the bait. No Hamadryas were found here on either of the two remaining 
baits or on G. ulmifolia fallen fruits nearby. 

Of the five female specimens of Hamadryas collected for examination of reproductive 
condition, only one individual of H. guatemalena had a single mature (sculpted surface) 
egg and all others either had no ova at all or immature ova, in spite of (a) appearing in 
relatively “fresh” wing condition and (b) with one exception, being mated as evidenced 
by the presence of single spermatophore (Table 2). The spermatophore found in the 


VOLUME 39, NUMBER 3 233 


TABLE 2. Evidence for a lack of female reproductive activity (egg production) in 
Hamadryas species in northwestern Costa Rica during the tropical dry season. 


Female 


Species* no. Condition of ovary Spermatophore** 

H. guatemalena 1 no ova one present, “fresh” (mated) 

guatemalena (Bates) 2 no ova one present, “fresh” (mated) 
3 one near full size one present, broken, (mated) 
egg only “old” 

H. februa ferentina 1 no ova none (unmated) 
(Godart) 

H. iphthime iphthime 1 four immature one present, “fresh” (mated) 
(Bates) ova 


en the exception of one female of H. guatemalena, all of these butterflies were collected at the “Barranca Site” 
oca ity 

** “Fresh” spermatophores appeared full and een peor: exuded a milky fluid, and were intact within the bursae 
copulatrix; “old” spermatophore appears collapsed and fragmented. 


individual with the single mature egg appeared “old” since it was easily fragmented 
during the dissection, whereas, other spermatophores appeared “‘fresh”’ (Table 2). 

Our data, while preliminary and based upon small sample size, does point out some 
interesting new information about the natural history of Hamadryas in lowland tropical 
dry forest during the dry season: (1) occurrence of individual species may vary consid- 
erably over relatively small distances (e.g., 40 km); (2) behavioral posturing associated 
with nocturnal perching is very different from that of daytime perching, both in the site 
of perching and the posture of wings; (3) the choice of both daytime and nocturnal 
perching sites may be determined in part by the location of trees and foliage in wind- 
sheltered places and having little or nothing to do with thermoregulation in response to 
dryness; (4) by the middle of the lengthy dry season there may be little or no reproductive 
activity as indicated by the absence of mature ova in most female specimens examined; 
(5) adults active at or near the middle of the dry season may represent the final “wave” 
of adults to eclose during this season as seen by their “fresh” wing condition and that 
these individuals do mate; and (6) there may be little or no adult feeding at this time, 
and little or no egg placement as well. Young (op. cit.) studied the life cycle of H. februa 
at the Barranca Site early into the dry season (December) and observed egg placement 
and successful larval development on Dalechampia, which was still evergreen at this 
time. He might have been studying the immature stages of a crop of fresh adults that 
would be active later in the dry season at this locality. Ehrlich and Ehrlich (1978, J. 
Kansas Entomol. Soc. 51:666-697) collected a single female of H. feronia which had 
thirteen ova, a figure in sharp contrast with the ova-less females found in our study. We 
interpret this difference to highlight the lack of reproductive activity in these butterflies, 
as seen in our sample, during the latter half of the tropical dry forest lengthy dry season. 
Spermatophores present in these butterflies at this time appeared fresh (i.e., filled with 
milky white fluid) and perhaps are used to fertilize ova that may develop at the end of 
the dry season and early into the rainy season. As we did not find larval food plants at 
the Liberia locality and did not check for them at the Barranca Site, we can only 
tentatively speculate that they were in short supply at this time of the year. We suggest 
that Hamadryas undergo a reproductive diapause in ovarian development during the 
latter half of the dry season and that the presence of fresh spermatophores at this time 
is an adaptation to facilitate egg maturation and egg placement when the rainy season 
begins. The observed absence of mating plugs suggests that these females may mate 
again, perhaps at the end of the dry season or early into the rainy season, if they survive. 
Butterflies thriving in open areas and at the edges of forest habitats in the tropics may 
necessarily experience such cessations in breeding in response to diminished resources 
for immature stages. The association of Hamadryas with such habitats is well known 
(e.g., Ross, 1976, op. cit.; Jenkins, 1988, op. cit.; Schwartz, 1983, Mus. Nac. Hist. Nat. 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Santo Domingo, 69 pp.). Adult butterfly populations in the seasonal tropics generally 
decline as the dry season advances, and there is less reproductive activity at this time, 
both in the New World and Old World tropics (e.g., Young, 1981, Oct. Oecol. Ecol. 
Gener. 2:17-30; Spitzer, 1983, J. Res. Lepid. 22:126-130). 

Declines in butterfly populations with the dry season may reflect contracting supplies 
of larval food (Spitzer, 1983, op. cit.), and reproductive activity may be restored as 
deciduous larval food plants produce tender, new meristems in the latter half of the dry 
season (Young, 1983, J. Lepid. Soc. 37:313-317). In one forest patch along the Pan 
American Highway between Canas and Liberia, we observed the moth Haemaorrhagia 
(Sphingidae) carefully placing eggs singly on the small (2-5 mm long) folded leaf mer- 
istems of an unidentified understory tree near dusk (2 March). Adults of many butterfly 
species were seen at the Barranca Site at this time, including: Morpho peleides Kollar, 
Caligo memnon Felder, Memphis morvus boisduvalo Comstock, Consul fabius Double- 
day, Siproeta stelenes Fruhst., Zaretis itys Cramer, Taygetis andromeda Cramer, Phi- 
laethria dido Linnaeus. Parides arcas mylotes Bates, Battus polydamas Linnaeus, and 
Papilio anchisiades idaeus Fabricius. Many plant species at this locality have small 
meristems at this time, and some butterfly species selectively oviposit on these tissues. 
For example, we observed Itaballia demophile calydonia Boisduval carefully placing 
eggs singly on very fresh leaf meristems of Capparis sp. (Capparidaceae) at 1530 h on 4 
March. Cuttings of meristem stem tissues of this plant species quickly wither, even when 
confined to tightly closed plastic bags, whereas, cuttings of older stems do not wither as 
fast. We interpret these observations to mean that considerable moisture stress is operative 
on butterfly food plants at this time and that new meristems may be a very limited 
resource for egg placement in various plant groups present. 

Our data suggest that Hamadryas adults devote considerable time daily to perching 
on tree trunks in sheltered places. Many insects living in moisture-stressed habitats care- 
fully position themselves to minimize direct exposure to sustained dryness and related 
ambient factors (e.g., Egwuatu, 1980, Z. Angew. Entomol. 90:347-354; Toms, 1981, Zool. 
Zh. (U.S.S.R.) 60:1486-1494; Shelly, 1982, Physiol. Zool. 55:335-348; Gillis & Possai, 1983, 
Ecol. Entomol. 8:155-161; Findlay et al., 1983, Ecol. Entomol. 8:145-153; Chappell, 
1983, Anim. Behav. 31:1088-1093; Shiffer, 1983, J. Med. Entomol. 20:365-370). Often- 
times, an insect species in a particular habitat will exhibit strong diurnal changes in 
distribution in response to day-night cycles of both temperature and illuminescence (e.g., 
Van Etten, 1982, Entomol. Exp. Appl. 32:38—45; Parker, 1982, Amer. Mid]. Nat. 107: 
228-237). The nymphalid butterfly Anartia fatima Fabricius forms loose “aggregations” 
of adults in wind-sheltered bushes for nocturnal perching during the Guanacaste dry 
season (Young, 1979, op. cit.). The observed tendency for adult Hamadryas to perch 
both day and night on the wind-sheltered edge of a forest patch may reflect a concen- 
tration of “nuclear” adult populations around such places during the dry season. Medi- 
terranean fruit flies are present in higher densities in traps in dry areas than in wet areas 
in the Hawaiian Islands (Vargas et al., 1983, Environ. Entomol. 12:303-310). The same 
species of insect may selectively choose different plant parts as perching sites at different 
times of the diurnal cycle, as witnessed in our study. Tsetse flies in Africa perch on woody 
plant parts in the day and on leaves at night (Turner, 1980, Insect Appl. Sci. 1:15-21). 

Our data clearly suggest a tenacity of Hamadryas to forest sites in the seasonal tropics, 
an adaptive response, we suggest, to increasing the survival of small populations of 
diapausing adults in protected places until the rainy season begins and larval food plants 
leaf out. Many Euphorbiaceae exhibit marked seasonal cycles in vegetative growth in 
the tropics (Lieberman, 1982, J. Ecol. 70:791-806). And while dry season weather con- 
ditions may have adverse effects on egg-laying activity in butterflies (e.g., Zalucki, Res. 
Popul. Ecol. 23:318-327), we suspect that severe larval food plant availability is the 
prime factor selecting for dry season diapause in adult Hamadryas during the later phases 
of the dry season in this region of Costa Rica. We do not extend these predictions to 
other, i.e., less seasonal, regions of Central America where these butterflies also occur. 
“Fresh” but mated female Hamadryas may exhibit a preference for staying in and near 
forest patches during the dry season and may move away from these sites in search of 


VOLUME 39, NUMBER 3 235 


oviposition sites when older. Older, mated tsetse flies exhibit different habitat preferences 
from new females in the Ivory Coast (e.g., Gouteux, 1982, Cah. Orstom. Ser. Entomol. 
Med. Parasitol. 20:41-61). In short, we might have missed finding older female Hama- 
dryas (with “worn” wings) since our census program was very limited. But, both “fresh” 
and “worn” males and females of Morpho peleides Kollar (Morphidae) exhibit confined 
movements in the Barranca Site forest throughout the dry season (Young & Thomason, 
1974, op. cit.). 

We were surprised to find no evidence of adult feeding in our brief study, with the 
exception of observing one adult on the banana bait at the Barranca Site. While these 
butterflies are known to feed on sweet smelling rotting fruits (e.g., Schwartz, 1983, op. 
cit.) as well as on tree sap (Ross, 1976, op. cit.) and horse dung (Jenkins, 1983, op. cit.), 
we did not observe feeding on naturally occurring food sources. The cattle trail that runs 
to one side of the trees along the highway had piles of fresh dung (cattle and horse). 
Jenkins (1988, op. cit.) suggests a preference for horse dung over cattle dung by these 
butterflies. Whether or not there is a cessation of adult feeding in the dry season physi- 
ologically linked to a probable reproductive diapause, remains to be studied. Subsequent 
to this study, one of us (A.M.Y.) observed H. amphinome mexicana (Lucas) feeding on 
freshly fallen “guava” fruits at “Finca La Tirimbina,” near La Virgen (10°23’N, 84°07'’W; 
220 m), Sarapiqui District, Costa Rica on 2—4 August 1984. Several butterflies were seen 
inserting their probosci into small wounds in the fruits (1100-1400 h). 

Because shaded pockets of forest in highly seasonal tropical localities are refugia for 
many animals during the dry season, predation upon adult Hamadryas may be exceed- 
ingly high at these times, further selecting for avoidance of feeding, particularly on the 
ground. Lizards and birds figure prominently as predators on adult Hamadryas (Jenkins, 
1983, op. cit.). Different types of predators, attacking different life stages, may act at 
different seasons to exploit Hamadryas populations. If predation on adults is high during 
the dry season, it might be lower in the rainy season and replaced at these times by 
increased mortality of immature stages. In the seasonal tropics, some insect populations 
are regulated’ by varying sets of mortality factors associated with seasonality (e.g., 
Page, 1980, Bull. Entomol. Res. 70:621-633). At times of the year when both larval and 
adult food resources are abundant, immature stages of Hamadryas populations may build 
up most intensely in the vicinity of adult resources, as suggested by some temperate zone 
butterfly studies (Murphy, 1983, Environ. Entomol. 12:463-466). When this occurs, 
Hamadryas caterpillars may become a major herbivore of Dalechampia situated near 
adult feeding sites (Armbruster, 1982, op. cit.). However, such an interaction is predicted, 
on the basis of our preliminary results, to be inoperative during the latter half of the dry 
season in the lowlands of northwestern Costa Rica. 

This study was made possible by a grant from The Friends of the Museum of the 
Milwaukee Public Museum. We thank Joan P. Jass for field assistance. 


ALLEN M. YOUNG AND SUSAN S. BoRKIN, Invertebrate Zoology Section, Milwaukee 
Public Museum, Milwaukee, Wisconsin 58288. 


Journal of the Lepidopterists’ Society 
39(3), 1985, 235-236 


DONATION OF BLANCHARD LEPIDOPTERA COLLECTION 
TO THE SMITHSONIAN INSTITUTION 


Until recently, few regions of comparable diversity in the United States had been as 
little surveyed for Lepidoptera as the State of Texas. Thus, it is with considerable grat- 
itude and respect that the Smithsonian Institution acknowledges the donation of the 
largest and finest prepared collection of Lepidoptera ever assembled from this region, as 
well as the total accomplishments of Mr. André Blanchard, the man responsible. 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The fascinating life of André Blanchard consists of a sequence of successful careers 
too diverse to summarize adequately in this short note. During his earliest career with 
the French Navy, he served in numerous capacities, ranging from a seaplane pilot to the 
commander of the research ship Les Eparges. Following a distinguished military career, 
he eventually became head of the Physics Laboratory of the Michelin Tire Company, 
transferring in 1943 to Schlumberger, where he rose to Vice President of Research and 
Development. Interspersed between these responsibilities was a brief career during World 
War II when Blanchard served as a translator for the War Department. 

Of most concern to entomologists were Blanchard’s activities following his retirement 
from Schlumberger in 1961. It was then that he began to survey the Lepidoptera, moths 
in particular, of his adopted state, Texas. He had previously collected Lepidoptera in his 
native France, but never with such dedication nor determination. His accomplishments 
since 1961 must surely be an inspiration to anyone contemplating retirement. Not content 
with amassing a fine and valuable collection, Blanchard quickly assumed an active re- 
search interest on the moths of Texas. Thus far, he has authored or co-authored 51 papers 
in this series, mostly treating the Pyralidae and Tortricidae. 

The Blanchard Collection totals 76,852 specimens of which 60,233 are Lepidoptera 
and 16,305 are Coleoptera. The Macrolepidoptera are the best represented, particularly 
the Noctuidae (over 18,500 specimens) and Geometridae (ca. 8000). The Pyraloidea (over 
9000) and Tortricoidea (ca. 4000), which became a major focus of his over the last decade, 
are also strongly represented. Some of the larger Microlepidoptera (e.g., Acrolophinae) 
are likewise present in large series. Included in the collection are 82 holotypes and over 
700 paratypes. The research value of the Blanchard Collection is further enhanced by 
more than 4600 microslides, mostly of genitalia. 

Mr. Blanchard continues to reside in Houston, Texas with his second wife May Elise, 
who was instrumental in assisting her husband in his collecting efforts. 


DONALD R. Davis, Depertment of Entomology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, D.C. 20560. 


VOLUME 39, NUMBER 3 237 


Journal of the Lepidopterists’ Society 
39(3), 1985, 237 


BOOK REVIEW 


THE LIFE HISTORIES OF THE BUTTERFLIES IN JAPAN, Volume II and III (1984), by H. 
Fukuda, E. Hama, T. Kuzuya, A. Takahahi, M. Takashi, B. Tanaka, M. Tanaka, M. 
Wakabayashi, and Y. Watanabe, Hoikusha Pub. Co. Ltd., Osaka, 540 Japan. 


These books are part of a series describing the life histories of Japanese butterflies. 
Volume II is devoted to the Nymphalidae and Libytheidae and Volume III to the Ly- 
caenidae. Both books are published in Japanese with English summaries. 

The books begin with a series of excellent color plates showing the adult, egg, larva, 
pupa, host plant and often the habitat for each butterfly species covered in the book. 

The text describes the distribution, habitat, food plants, flight period, and early stages 
of each species found in Japan, including common migrants. Next comes the English 
summary of the text which covers the same material in less detail. The text is referenced 
to the plates by the name of the butterfly and plate number. 

There are distribution maps and an index to generic and specific names, both in 
English. The Table of Contents is in English and Japanese and gives the plate number 
and text location for each butterfly. 

These books are well bound and have two cloth bookmarks attached to the spine. The 
front covers have an excellent color photograph of a native butterfly. Volume II costs 
4500 yen and Volume II 5000 yen. 

I found the English summaries sparse, and it was obvious that more data are contained 
in the Japanese text. However, each summary contained a complete capsule of infor- 
mation about the butterfly. The pictures are excellent and show the butterflies in their 
habitats and on their food plant. 

These books are obviously aimed at a Japanese reading audience. However, the English 
summaries and plates make them a welcome addition to the library of any butterfly 
collector. 


ROBERT V. DOWELL, Analysis and Identification, California Department of Food 
and Agriculture, 1220 N Street, Sacramento, California 95814. 


Journal of the Lepidopterists’ Society 
89(3), 1985, 238 


EUREMA NISE IN JAMAICA 


One of the most fascinating accounts of butterfly rediscovery in recent years is that of 
Eurema nise nise in Jamaica (Klots & Heineman, 1957, Proc. R. Entomol. Soc. Lond. 
(B)26:206-214, Plate I). E. nise, originally described and figured from Jamaica by Cramer 
in 1775, eluded collectors in Jamaica until a young enthusiast, G. Irving Latz, accom- 
panying experienced lepidopterist Bernard Heineman, netted one in March 1951 in St. 
Ann Parish. In the 175 years between the description and the capture by Latz, there was 
much discussion in lepidopterological literature regarding the true origin or identity of 
Cramer’s nise. During that time, also, nise was widely and commonly found in other 
Antillean Islands and on the continent from the extreme southern USA to Uruguay and 
mid-Argentina and was described under various names, a number of which are consid- 
ered today as valid subspecies. 

Since 1951, a quantity of nise specimens have been collected in Jamaica. Riley in his 
popular guide (1975, A field guide to the butterflies of the West Indies, p. 120) states, 
“Reported only in January and February, i.e. the winter brood, but a summer brood 
must also occur.” This, however, is not true, since Klots and Heineman (1957) and Brown 
and Heineman (1972, Jamaica and its butterflies) cite known specimens from Jamaica 
dated June, July, August and September, some of which are figured and are clearly of 
the summer phenotype. What is true, is that the great majority of the specimens collected 
there were found in the winter months (when most collectors visit Jamaica) and are of 
the winter phenotype. 

I have had many years’ experience with, and have handled hundreds of specimens of, 
Eurema nise tenella (Boisduval) 1836 from northwestern Argentina. There, the winter 
phenotypes are affected in two ways: (1) There is a progressive reduction of the upperside 
black borders, and (2) there is an augmentation of the rusty-brown scaling and blotching 
on the ventral hindwings. These same phenomena also take place in nearly all the other 
species of Eurema, as well as many other coliadine Pieridae flying in Argentina. What 
is of special interest is that these two phenomena do not necessarily occur simultaneously, 
which results in a number of distinct winter phenotypes. The literature in recent years 
has contained many opinions and a number of serious studies (some of which are con- 
tradictory) as to the causes of these phenomena in various species of the Coliadinae. 
Decrease in temperature, decrease in humidity, decrease in photoperiod or a combination 
of two or more of these factors during the immature stages have all been suggested as 
the cause. It is not the purpose of this note to add another field-based opinion to the 
confusion. Whatever the causes, the result is the great diversity of forms of E. nise both 
in Jamaica and Argentina, as are in part illustrated in the works mentioned above. 

This note reports the collection of 14 specimens of E. nise nise in St. Andrew, Trelawny 
and Manchester parishes of Jamaica between 22 and 26 November 1983 and that these 
are basically of what I call the “autumn” phenotype, comparable to specimens of E. nise 
tenella captured in lowland northwestern Argentina in the month of May. This pheno- 
type is very close to the summer one, differing by having a very slight reduction of the 
dorsal wing borders, but like the summer phenotype showing no or very minimal rusty- 
brown scaling on the ventral hindwing. Voucher specimens from Jamaica have been 
deposited in the Allen Museum of Entomology and the National Museum of Natural 
History (Smithsonian). 

The specimens taken in St. Andrew Parish constitute a new parish record. Other 
Eurema collected in Jamaica during November 1983 were E. daira palmira, E. messa- 
lina, E. nicippe (St. Andrew Parish—record), E. adamsi, E. proterpia, E. lisa euterpe 
and E. dina parvumbra. A number of these are considered in the literature to be scarce. 
Most were seen, though not captured, in quantities. This would indicate that November 
is an exceptionally good month for Eurema in Jamaica. 

I thank Dr. Lee D. Miller and Dr. Robert K. Robbins for their helpful criticisms of 
the manuscript. 


ROBERT C. EISELE, Casilla de Correo 90, 4107 Yerba Buena (Tecuman), Argentina. 


Date of Issue (Vol. 39, No. 3): 13 May 1986 


EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


THomas D. EICHLIN, Retiring Editor 


Associate Editors: 
BoyYcE A. DRUMMOND III, DOUGLAS C. FERGUSON, THEODORE BD. SARGENT 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
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plant or animal in the text should include the full scientific name, with authors of 
zoological names. Insect measurements should be given in metric units; times should be 
given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where 
italics are intended. References to footnotes should be numbered consecutively, and the 
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Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


A NEw SPECIES OF TILDENIA FROM ILLINOIS (GELECHIIDAE). 
Ronald W. Hodges. Ee 
OBSERVATIONS ON THE BIOLOGY OF PARNASSIUS CLODIUS (PAPIL- 
IONIDAE) IN THE PACIFIC NORTHWEST. David V. McCorkle 
d> Paul C. Hammond ou 
THE BIOLOGY AND IMMATURE STAGES OF AUTOMERIS RANDA AND 
AUTOMERIS IRIS HESSELORUM (SATURNIIDAE). Paul M. 
Tuskes) co 


COURTSHIP AND OVIPOSITION PATTERNS OF TWO AGATHYMUS 
(MEGATHYMIDAE). Don B. Stallings, Viola N. T. Stallings, 
J. R. Turner & Beulah R. Turner _.._.. 


BIOLOGY OF THE HALF-WING GEOMETER, PHIGALIA TITEA CRA- 
MER (GEOMETRIDAE), AS A MEMBER OF A LOOPER COMPLEX 
IN WEST VIRGINIA. Linda Butler... ee 


THE RELATIONSHIP BETWEEN PEDALIODES PERPERNA AND PE- 
TRONIUS (SATYRIDAE), WITH THE DESCRIPTION OF A NEW 
SUBSPECIES. Lee D. Miller’ 

ECOLOGICAL NOTES ON SYNANTHEDON DOMINICKI DUCKWORTH 
AND EICHLIN (SESIIDAE) IN FLORIDA AND FIRST DESCRIP- 
TION OF THE FEMALE. Larry N. Brown, Thomas D. Eich- 
lin.d> J. WendellSnow 0 

How TO DO GENETICS WITHOUT MAKING THE BUTTERFLIES 
Cross.” John R.G. Turner 20 

NOTES ON PSEUDOSPHINX TETRIO (L.) (SPHINGIDAE) IN PUERTO 
Rico. Jorge A. Santiago-Blay _...... 


GENERAL NOTES 
Natural History Notes on Astraptes and Urbanus (Hesperiidae) in Costa Rica. 
Allen M, Young Sich 


Independent Evolution of “False Head’ Behavior in Riodinidae. Robert K. 
Robbins. ys ei sie ie a 


Interactions of Parasitoids with an Opsiphanes (Brassolidae) Caterpillar in 
Costa Rica... Allen M. Young 2200 


On a Previous Report of Diurnal Roosting of the Pipevine Swallowtail, Battus 
philenor (L.):. Raymond'W, Neck 220000 


Natural History Notes for Some Hamadryas Butterflies (Nymphalidae: Nym- 
phalinae; Aegeronini) in Northwestern Costa Rica during the Tropical 
Dry Season. Allen M. Young ¢> Susan S. Borkite ccc cece ceeeeeteteeeeeeoee 


Donation of Blanchard Lepidoptera Collection to the Smithsonian Institution. 
Donald BR. Davis: iio es a i 


Eurema nise in Jamaica. Robert C. Eisele 
Book REVIEW 


Volume 39 1985 Number 4 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





25 June 1986 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CLIFFORD D. FERRIS, President DOUGLAS C, FERGUSON, 

Don R. Davis, Immediate Past President President-Elect 

JERRY A. POWELL, Vice President EDWARD M. PIKE, Vice President 
RICHARD A. ARNOLD, Secretary ALLAN WATSON, Vice President 


ERIC H. METZLER, Treasurer 


Members at large: 


JOHN M. BuRNS Boyce A. DRUMMOND III MIRNA. M. CASAGRANDE 
FLOYD W. PRESTON JOHN LANE EDWARD C. KNUDSON 
JACQUELINE Y. MILLER ROBERT K. ROBBINS FREDERICK W. STEHR 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229, U.S.A.; and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$25.00 (institutional subscriptions) and $18.00 (active member rate) by the Lepidopter- 
ists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Boule- 
vard, Los Angeles, CA 90007. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, 
1900 John St., Manhattan Beach, CA 90266. 


Cover illustration: Micropylar end view (x 130) of the egg of Sericosema sp. (probably 
juturnaria) (Geometridae). The scanning electronmicrograph was taken by Thomas D. 
Eichlin, Sacramento, of eggs furnished by Ron Robertson, Santa Rosa, California. 


JouRNAL OF 
Tue LEPIDOPTERISTS’ SOCIETY 


Volume 39 1985 Number 4 


Journal of the Lepidopterists’ Society 
39(4), 1985, 239-261 


BIRD PREDATION ON LEPIDOPTERA AND THE 
RELIABILITY OF BEAK-MARKS IN 
DETERMINING PREDATION PRESSURE 


MARK K. WOURMS AND FRED E.. WASSERMAN 


Department of Biology, Boston University, 
Boston, Massachusetts 02215 


ABSTRACT. Visually hunting predators such as birds are thought to have influenced 
the evolution of the wing markings and colorations of Lepidoptera. Although studies 
have been conducted to quantify and characterize predation by birds on butterfly pop- 
ulations, field observations of bird predation on butterflies have rarely been reported. A 
request for information on predation yielded 50 previously unpublished accounts of bird 
predation on butterflies. 

The combination of laboratory interactions of Pieris rapae and blue jays and field 
collections of P. rapae allowed several variables to be examined which affect the reli- 
ability of using frequency of beak-marks on lepidopteran wings as an index of predation 
pressure. Beak marks occur four times more frequently during attacks on flying P. rapae 
than on ones at rest and blue jays were five times more efficient at capturing resting 
butterflies than capturing flying butterflies. Variation in wing strength makes the area 
where the ipsilateral wings overlap and the costal vein area of the forewing more resistant 
to beak-marks than the marginal areas of the fore- and hindwings and the distal tip of 
the forewings. These differences in wing strength may confound the use of beak-marks 
as an index of predation pressure. 

Finally, predation efficiency and the frequency of occurrence of beak-marks during 
attacks, as determined in the laboratory, were used in conjunction with field data to 
estimate avian predation pressure on P. rapae populations. 


Although birds have long been thought to be the major predators on 
adult Lepidoptera (Poulton, 1890, 1913; Fryer, 1913; Swynnerton, 1915; 
Dover, 1920; Carpenter, 1937), field observations of bird predation on 
butterflies in temperate North America have rarely been reported. The 
short time that it takes birds to capture and manipulate butterflies 
while feeding may account for the rarity of field observations (Bowers 
& Wiernasz, 1979; Collins & Watson, 1983). There is strong circum- 
stantial evidence in the form of beak-marks and tears on wings of 
Lepidoptera to indicate that birds act as significant predators on but- 
terflies (e.g., Wheeler, 1935; Carpenter, 1937; Kolyer, 1968). 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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VOLUME 39, NUMBER 4 





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JOURNAL OF THE LEPIDOPTERISTS’ 


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JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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245 


VOLUME 39, NUMBER 4 





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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


246 


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247 


VOLUME 39, NUMBER 4 


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248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Predation by birds on Lepidoptera have been reported in studies 
that have been concerned with interactions between European Lepi- 
doptera and birds (Carpenter, 1933, 1937, 1941; Collenette, 1935) or 
tropical Lepidoptera and birds (Fryer, 1913; Young, 1971; Brown & 
Neto, 1976; Smith, 1979; Collins & Watson, 1983). This study reports 
on the interactions between North American Lepidoptera and birds 
and investigates the reliability of butterfly wing-damage frequencies 
as a predictor of predation pressure in the European cabbage butterfly, 
Pieris rapae L. 


METHODS 


A request for information from professional and amateur lepidop- 
terists and ornithologists regarding butterfly-bird interactions yielded 
50 previously unpublished accounts of predation by birds on butterflies 
in temperate North America (Table 1). The results of a literature sur- 
vey of the frequency of beak-marks reported in butterfly populations 
is summarized in Table 2 and a survey of defensive compounds found 
in adult Lepidoptera is presented in Table 38. 

In order to document avian predation on P. rapae in the field P. 
rapae adults were collected for a 30 minute period every seven to 10 
days in Boston, Suffolk Co., Massachusetts (Fenway Victory Gardens) 
and for a one hour period every seven to 10 days at two sites in 
Lexington, Middlesex Co., Massachusetts (Dunback Meadows and Car- 
roll Field, 71° West, 42° North). The difficulty of moving through the 
Middlesex Co. sites, due to dense vegetation, Phragmites spp. and 
goldenrods, Solidago spp., necessitated the longer collection time per 
period. Captured butterflies were sexed, and the presence and location 
of bird-attributable wing damage were recorded for each specimen. 
Initially, nine possible locations of attack were identified. These were 
condensed to represent three directions of attack; from the front, side, 
or from behind (Fig. 1). 

One factor that may influence the reliability of beak-marks as an 
index of predation is the strength of the wings. The strengths of (1) 
three areas on the forewing, (2) one area on the hindwing, and (3) the 
area where the ipsilateral fore- and hindwing overlap, were measured 
on 25 specimens of P. rapae (Fig. 2). Strength measurements were 
obtained by removing the wings from the specimen, and positioning 
one wing at a time in the testing device (Fig. 3). The device slowly 
increased the force on the wing until tearing occurred. Data were 
analyzed with a single factor repeated measures analysis of variance 
and a Student Newman-Keuls multiple pairwise test (Zar, 1974). 

To observe predatory behavior and to quantify the frequency and 


VOLUME 39, NUMBER 4 


249 


TABLE 2. Frequency of bird-attributable damage on the wings of Lepidoptera. 


Lepidoptera 


Colias eurytheme 
Pieris rapae 

P. rapae 

Pieris protodice 
Pieris coenia 


Ascia monuste 


Lycaenid spp. 


Lycaenid spp. 
Lycaenid spp. 
Euphydryas chalcedona 


Danaus plexippus 


D. plexippus 
D. plexippus 


Danaus chrysippus 

Hypolimnas misippus 

Morpho amathonte 
centralis 

Morpho granadensis 
polybaptus 

Morpho peleides 
limpida 

Cercyonis pegalia 


Maniola jurtina L. 


Catocala spp. 


Family 


Pieridae 
Pieridae 
Pieridae 
Pieridae 
Pieridae 
Pieridae 
Lycaenids 
Hairstreaks 
Lycaenids 


Lycaenids 
Nymphalidae 


Nymphalidae 


Nymphalidae 
Nymphalidae 
Nymphalidae 
Nymphalidae 
Morphidae 
Morphidae 


Morphidae 


Satyridae 


Satyridae 


Noctuidae 


Frequency of 
bird damage 


4.8% 


5.1% 


7.9% 
9.9% 


6.8% 


6.8% 


22.8% 


10% 
7.9% 
7.9% 
7.0% 


5.4% 
8.1% 


2% 


40% 


30.7% 


7.3% 
3.2% 
0% 


65.3% 
83% 


83% 


10% 
7.1% 


males 
females 


males 
femlaes 


8% males 
13% females 


4% 


Comments and references 


Sacramento Valley, Cali- 
fornia, Shapiro (1974) 

Sacramento Valley, Cali- 
fornia, Shapiro (1974) 

Boston, Massachusetts, 
Wourms (this study) 

Sacramento Valley, Cali- 
fornia, Shapiro (1974) 

Sacramento Valley, Cali- 
fornia, Shapiro (1974) 

Everglades Nat. Park, 
Florida, Pought and 
Brower (1977) 


Malaya, Robbins (1978) 

Colombia, Robbins (1978) 

Colombia, Robbins (1978) 

Panama, Robbins (1978) 

San Mateo Co., Califor- 
nia, Bowers, Brown 
and Wheye, submitted, 
1983 

Santa Cruz, Mexico, 
Tuskes and Brower 
(1978) 

Mexico, Calvert et al. 
(1979) 

Mexico, Carpenter and 
Hope (1941) 

Tanzania, Smith (1979) 

Tanzania, Smith (1979) 

Costa Rica, Young (1971) 


Costa Rica, three loca- 
tions, Young (1971) 

Costa Rica, three loca- 
tions, Young (1971) 

Massachusetts, two sites, 


Bowers and Wiernasz 
(1979) 


Southern Sweden, Bengs- 
ton (1981) 


Massachusetts, Sargent 
(1973) 





JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


250 


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VOLUME 39, NUMBER 4 251 


FRONT 
MIDDLE 


BEHIND 


Fic. 1. Front, middle, behind locations of bird damage for field collected Pieris 
rapae. 


type of butterfly wing damage that occurs during attacks, P. rapae 
adults were brought into the laboratory in a wire cage (25 cm high x 
15 cm in diameter), where they were released into a 1 x 0.5 x 1 m 
holding cage made of mosquito netting. Sugar water and wild flowers 
were provided ad libitum. Four blue jays, Cyanocitta cristata, were 
captured in mist nets and baited traps. They were housed individually 
in 1 x 1 X 1 m wire screen cages under a long-day light cycle (18 h 
light, 6 h dark). All birds were provided water and sunflower seeds ad 
libitum, and were given canned dog food, fresh chopped vegetables, 
and 5-10 mealworms each morning. Two weeks prior to trials with 
live P. rapae, one bird was placed in a flight cage (8 x 4 x 3 m). The 
experimental procedure consisted of (1) placing a single live P. rapae 
in a 4 cm box, (2) introducing the box into a flight cage through a slot 
in the side of the cage, and (3) releasing the butterfly by pulling a 
string attached to the lid of the box. 

The activities of the butterfly and the blue jay were monitored for 
15 minutes with a video recorder. If the butterfly was not consumed 
during the 15 minute trial it was removed and another individual was 
presented after a 15 minute interval. No more than six trials were 
conducted per day. Video tapes were analyzed with slow motion and 
freeze-frame to identify attacks and contact points. A new blue jay 
was transferred to the flight cage and trained, after the previous bird 
had had 10 days of live presentations. 


252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pel 


i a 
A. 
Forewing 
* 
pe 
B. 
* 
Hindwing 
Pe 


* 


Fic. 2. The areas of Pieris rapae wings where resistance to tearing was measured. 
(* indicates points of anchoring during measurements.) A. The arrows indicate points of 
strength measurements, the forewing costal vein, wing tip, and distal margin. B. Distal 
margin of the hindwing. C. Ipsilateral overlap of fore and hindwings. 


VOLUME 39, NUMBER 4 253 


lem 





tas 





23cm pe EB iy 


Fic. 3. Wing strength (resistance to tear) measurement device. The wing was an- 
chored by the clip of the Pesola scale, and the area measured by the lower clip. The 
crank was turned pulling upward. The resistance (g) was shown on the Pesola scale, and 
was recorded to the nearest 0.5 g the moment the wing tore apart. 


RESULTS 


Of the 1179 P. rapae collected during the three field seasons, an 
average of 7.2 + 0.28% (S.D.) of the specimens had beak-marks or 
beak tears. There were no significant differences in frequency of bird 
damage among sites or within sites over different seasons (Table 4, 6 x 
2 Chi-square contingency table; x? = 0.49, df = 4, P > 0.05). Only two 
of the 91 specimens collected showing bird damage had impressions 
of a bird’s beak on the wings of the butterfly; the other 89 specimens 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Butterfly sampling and beak-mark frequencies. 


Year Site Collected Beak-marked % damaged 

1981 Fenway 182 12 6.2 

1982 Fenway 351 28 7.4 
Lexington 104 8 7.1 

1983 Fenway 241 ily 6.6 
Lexington 247 22 8.2 
Carroll o4 A 69 

Totals IITA) 91 (iors as (0) 746) 


had beak tears. Henceforth, unless otherwise stated, ““beak-marks”’ im- 
plies both marks and tears. 

In 1981, sex was not distinguished during the collection of P. rapae. 
In 1982 and 1983, 838 (78.2%) males and 238 (21.8%) females were 
collected (Table 5). Chi-square analysis reveals that in 1982 and 1983 
the frequency of bird damage on P. rapae was independent of sex 
(Table 5). 

No specimen showed evidence of more than one attack. Symmetrical 
damage on both sets of wings suggests that the damage occurred while 
the butterfly was at rest with wings folded. Butterflies with damage on 
one wing or on an ipsilateral forewing and hindwing were assumed to 
have been attacked in flight (Bowers & Wiernasz, 1979; Sargent, 1978). 
Two specimens from 1982 and one from 1983 were omitted because 
symmetrical or single wing damage could not be determined. 

Of the 76 bird-damaged P. rapae collected in 1982 and 1983, 51 
(67%) were damaged in flight, and 25 (83%) were damaged at rest 
(Table 6). Significantly more specimens were damaged in flight than 
at rest (expected values are calculated as half of the total number of 
damaged specimens; x? = 4.4, df = 1, P < 0.05). The distributions of 
attacks from the front, side and from behind are presented in Table 
6. There was no significant difference between the distribution of at- 
tack positions occurring in flight from the distribution of attack posi- 
tions occurring at rest (Table 6; 3 x 2 Chi-square contingency table; 
x? = 2.44, df = 2, P > 0.05). Regardless of whether the butterfly was 
in flight or at rest, significantly more bird damage occurred from be- 
hind than from the side or from the front (expected values are calcu- 
lated assuming equal numbers from each of the three directions: Table 
6; flight, x? = 25.52, df = 2, P < 0.05; rest, x? = 18.89, df = 2, P < 0.05). 

Analysis of variance indicated a significant difference in strengths 
of the five wing areas (F = 68.3, df = 96, P < 0.05). The costal vein 
area and the ipsilateral overlap were three times stronger than the 
distal margin of the hindwing and twice as strong as the distal margin 


VOLUME 39, NUMBER 4 2a 


TABLE 5. Comparison of the sex ratios of bird damaged and undamaged specimens 
for each year. 


Damaged Undamaged Total 

1982 
Male 27 345 372 
Female 9 110 119 
Total 36 455 491 

x? = 0.012, df = 1, P > 0.05 

1983 
Male 32 434 466 
Female Ac 108 EIS 
Total 43 542 585 


x? = 0.82, df = 1, P > 0.05 


of the forewing (Table 7). Using Student-Newman-Keuls multiple pair- 
wise test, we found no significant difference between the costal vein 
area and the ipsilateral overlap area, and no significant difference be- 
tween the margins of the fore- and hindwings and the tip of the fore- 
wing, but there was a significant difference between these two groups 
of wing areas (P < 0.01). 

The presentation of 104 P. rapae to four blue jays in a flight cage 
resulted in 182 attacks and 69 butterflies captured (Table 8). Sixty- 
nine percent (57/83) of the attacks on resting butterflies resulted in 
captures, while only 12% (12/99) of the attacks on flying P. rapae 
resulted in captures. The blue jays were significantly more efficient in 
capturing butterflies at rest than in flight. (Table 8; x? = 27.78, df = 1, 
P < 0.05). 

Few of the butterflies that were attacked showed wing damage. Of 
the 83 butterflies attacked at rest, only one of the 21 P. rapae which 
escaped had wing damage. Only four of the 87 P. rapae which escaped 
attacks in flight received wing damage. 


DISCUSSION 


Despite the presence of mustard oils (Rothschild et al., 1970; Aplin 
et al., 1975) P. rapae were acceptable prey to blue jays in the field and 
laboratory and to house sparrows, Passer domesticus, purple martins, 
Progne subis subis, and various other avian species in the field (Table 
1). In this study, an average of 7.2% of the P. rapae collected showed 
evidence of attacks by birds in the form of beak imprints and beak 
tears, and no specimen showed evidence of being attacked more than 
once. In California, Shapiro (1974) collected P. rapae and found 5.6% 
of the specimens bird-damaged, and between 0.383% and 0.50% of the 


256 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 6. Frontal and rear attacks on P. rapae. 


Attacked in flight from Attacked at rest from 
Year Site Behind Side Front Behind Side Front 
1983 Fenway 8 3 0 3 0 2 
Lexington 8 1 4 6 ] 2 
Carroll 0 1 1 2 0 0 
1982 Fenway 14 Zz 4 3) 0 2 
Lexington A al 0 mz 0 0 
Total 34 8 9 18 ] 6 


damaged specimens had multiple beak-marks. The percentages of bird 
damage reported by Shapiro (1974) and this study fall within the range 
of damage found in other lepidopteran species studied (Table 2). 

At least four variables can affect the relationship between beak- 
marks and predation pressure (Benson, 1972; Shapiro, 1974; Robbins, 
1980, 1981): 1) Damage may occur more readily during attacks on 
flying Lepidoptera than during attacks on resting Lepidoptera; 2) dif- 
ferent avian predators may be many times more successful during 
attacks on resting prey than during attacks on flying prey; 3) different 
avian predators may vary significantly in capture efficiency on Lepi- 
doptera; and 4) the strength of various butterfly wing areas differs, and 
this may influence the probability of obtaining beak-marks. 

The live presentations of cabbage butterflies to blue jays indicates 
that bird damage may occur up to four times more readily during 
attacks on flying butterflies than during attacks on resting butterflies. 
Therefore, if equal numbers of butterflies are attacked in flight and at 
rest, a field sample would reveal a greater number of specimens show- 
ing evidence of being attacked in flight due to the higher frequency 
at which damage occurs (Table 6). This was also found by Bowers and 
Wiernasz (1979) in C. pegala and is expected if avian predators are 
less efficient during attacks in flight than at rest. The reliability of a 
beak-mark predation index is seriously jeopardized by unequal chances 
of obtaining beak-marks in flight and at rest. If most predation occurs 
while the butterflies are in the vegetation and few beak-marks result, 
the index would underestimate the amount of predation occurring. 
Likewise, if most attacks occur in flight the amount of damage may 
be overestimated if many prey are damaged and few captured. No 
previous study had quantified the relative occurrence of beak-marks 
due to attacks at rest and in flight. 

Live trials support the hypothesis that avian predators may be much 
more efficient at capturing resting butterflies than at capturing butter- 
flies in flight (Table 8). The attack efficiency of blue jays on flying P. 


VOLUME 39, NUMBER 4 257 


TABLE 7. The strengths of P. rapae wing areas. 


Area Strength (g + S.D.) 
Forewing tip 9.24 + 2.4 
Forewing margin 8.28 + 2.62 
Hindwing margin 6.95 + 2.5 
Costal vein 19.76 + 6.0° 
Overlap of ipsilateral wings 20.16 + 4.2° 


The strength of wing areas with identical superscripts were not significantly different from each other. Groups with 
the ‘a’ superscript were significantly different from those with ‘b’ at the P < 0.01 level. 


rapae was 12%. This was lower than that of an aerial insectivore, the 
spotted flycatcher, Muscicapa striata, which was reported in the field 
to have captured four flying P. rapae in 17 attempts for a success rate 
of 23.5% (Davies, 1977) and was still lower than the success rate of 
100% on flying butterflies reported for a hunting northern shrike, La- 
nius borealis (Morrison, 1980; and pers. comm.). The predation index 
could be complicated by differences in the composition of avian com- 
munities in different habitats. For example, if shrikes were common 
in one habitat and relatively rare in another, many butterflies could 
have been consumed in the first area with little damage occurring, 
while the second area might have shown a high frequency of beak- 
marks but with little actual predation occurring. In this study, the avian 
communities of all three field sites were predominated by house spar- 
rows, Passer domesticus, song sparrows, Melospiza melodia, and Eu- 
ropean starlings, Sturnus vulgaris. 

Avian predators not only show variation in their probability of at- 
tacking butterflies, but may preferentially attack different areas on the 
butterfly in response to butterfly wing-markings. Butterflies attacked 
in stronger wing areas may show fewer beak-marks if they escape. 
Therefore, due to species differences in strengths of the areas of the 
wings, the frequencies of beak-marks may not be a reliable index of 
predation pressure for comparisons among species. 

The percentage of bird-damaged specimens actually represents only 
the number of individuals which successfully survived attacks and es- 
caped with bird damage. No data exist from field observations on the 
percentage of escaped butterflies showing no bird damage or on the 
percentages of attacked butterflies actually killed or eaten. If the lab- 
oratory efficiencies of blue jays preying on P. rapae are extrapolated 
to the field, avian predation on Lepidoptera becomes a much more 
significant selective force than previously suspected. Of the 1179 P. 
rapae collected, 76 had wing damage. Fifty-one specimens were at- 
tacked in flight, and 25 specimens were attacked at rest. In the labo- 
ratory only 4% of butterflies attacked in flight actually showed wing 


258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 8. Live presentations of P. rapae to blue jays. 


Presented Attacked Captured Efficiency (%) 
Bird Rest Flight Rest Flight Rest Flight Rest Flight 
I 29 11 41 22 28 3 68.3 13.6 
2 23 0 22 25 16 4 Uo 16.0 
3 It 12 18 41 11 tS) 61.1 12.2 
40g See Ae Zwei am, liste 100 0 
68 36 83 es) o7 12 68.7 12.1 


damage. Therefore, given the 4% probability of obtaining a beak-mark, 
the 51 specimens collected in the field that were attacked in flight may 
represent attacks in flight on approximately 1275 individuals. 

Blue jays consumed 12% of the P. rapae they attacked in flight in 
the laboratory. If this efficiency is extrapolated to the field, 12% of 
approximately 1275 P. rapae attacked or 1538 would have been con- 
sumed after capture in flight. 

Blue jays damaged only 1% of the butterflies attacked at rest in the 
laboratory. The 25 field-collected specimens which had damage from 
attacks while at rest would represent 2500 butterflies attacked at rest 
in the field. However, 68% of the resting P. rapae attacked in the 
laboratory were consumed. Therefore, according to this extrapolation, 
approximately 1700 P. rapae would have been consumed in the field 
after capture while at rest. 

The disparity in predation pressure in flight and at rest suggests that 
the major selective force of avian predation is directed at the butterfly 
wing surface that is exposed while the butterfly is at rest. This is the 
ventral surface of the wings for most butterflies (Platt et al., 1971) but 
may be the dorsal surface of the wings of most moths (Sargent & 
Keiper, 1969; Endler, 1978) and for butterflies which expose the dorsal 
surfaces of the wings during basking, nectaring, and other activities. 
Rawlins and Lederhouse (1978) found that in the Battus philenor 
mimicry complex, the resemblance of model and mimic is closest on 
the ventral wing surface. They suggest that selection may be most 
intense on the underside of the wings, which are exposed while the 
butterflies are at rest, rather than on the dorsal surface of the wings 
which is only exposed in flight. This hypothesis previously had not 
been evaluated critically, and in fact, is not supported by accounts of 
avian predation on Lepidoptera (Table 1). Attacks on resting butterflies 
may be less noticeable than attacks in flight because they occur rapidly 
and are often obscured by vegetation. 

Laboratory data obtained in the present study on predation by blue 
jays on P. rapae are the first to quantify differential predation pressure 


VOLUME 39, NUMBER 4 259 


on the ventral and dorsal surfaces of the wings of P. rapae due to 
variation in success rates of attacks on flying and resting butterflies. 
Recently, this has been supported by work on Euphydryas chalcedona 
(Bowers et al., unpubl. manuscript). Euphydryas chalcedona males 
which had less red on the dorsal surface of their wings were under 
greater predation pressure when their wings were spread while resting, 
basking, and nectaring. Although the ventral surfaces of the wings were 
essentially identical in both groups, avian predation appears to favor 
dorsally red males. Realistic estimates of predation pressure on Lepi- 
dopteran populations are impossible due to the lack of field data. Yet, 
the extrapolation of laboratory and field data supports the concept that 
bird predation on butterflies may be a more significant selective force 
on Lepidopteran populations than previously assumed. 

Many variables can influence the reliability of using the frequency 
of beak-marks on the wings of butterflies as an index of predation 
pressure. Thus, the interpretation of beak-mark frequencies is compli- 
cated and may not provide a reliable index of the amount of avian 
predation pressure on Lepidoptera. 


ACKNOWLEDGMENTS 


Financial support for these investigations was provided by the Frank M. Chapman 
Memorial research grant from the American Museum of Natural History, the Louis 
Agassiz Fuertes research grant, and the Boston University chapter of Sigma Xi. Thanks 
are also extended to the following people for help of various kinds; S. Duncan, T. Kunz, 
D. Phillips, R. Regis, B. Schlinger, and J. Traniello. 


LITERATURE CITED 


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white and small white butterflies (Pieris spp.) for the presence of mustard oils and 
mustard oil glycosides. J. Entomol. (A) 50(2):73-78. 

BENGTSON, S. A. 1981. Does bird predation influence the spot-number variation in 
Maniola jurtina (Lepidoptera)? Biol. J. Linnean Soc. 15:23-27. 

BENSON, W. W. 1972. Natural selection for Mullerian mimicry in Heliconius erato in 
Costa Rica. Science 176:936-938. 

Bowers, M. D. 1980. Unpalatability as a defense strategy of Euphydryas phaeton 
(Lepidoptera: Nymphalidae). Evolution 34(3):586-—600. 

Bow Ers, M. D. & D. C. WIERNASZ. 1979. Avian predation on the palatable butterfly, 
Cercyonis pegala (Satyridae). Ecol. Entomol. 4:205-209. 

BROWER, L. P., W. N. RYERSON, L. L. COPPINGER & S. C. GLAZIER. 1968. Ecological 
chemistry and the palatability spectrum. Science 161:1349-1350. 

Brown, K. S. & J. V. NETO. 1976. Predation on aposematic Ithomiine butterflies by 
tanagers. Biotropica 8(2):136-141. 

CALVERT, W. H., L. E. HENDRICK & L. P. BROWER. 1979. Mortality of the Monarch 
butterfly (Danaus plexippus): Avian predation at five over wintering sites in Mexico. 
Science 204:847-851. 

CARPENTER, G. D. H. 1983. Attacks of birds on butterflies. Trans. Entomol. Soc. Lond. 
81:21-26. 

1937. Further evidence that birds do attack and eat butterflies. Proc. Zool. Soc. 

Lond. (A) 107:2238-247. 





260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1941. An interesting sidelight on the causes of coloration in butterflies. Nature 
147:356. 

CARPENTER, G. D. H. & D. M. Hope. 1941. The relative frequency of beak-marks on 
butterflies of different edibility to birds. Proc. Zool. Soc. Lond. (A) 111:223-231. 
COLLENETTE, C. L. 1985. Notes concerning attacks by British birds on butterflies. Proc. 

Zool. Soc. Lond. 193:200-217. 

COLLINS, C. T. & A. WATSON. 1988. Field observations of bird predation on neotropical 
moths. Biotropica 15(1):53-60. 

Davies, N. B. 1977. Prey selection and the search strategy of the spotted flycatcher 
(Muscicapa striata): A field study on optimal foraging. Anim. Behav. 25:1016-1083. 

DONAHUE, J. P. & J. H. NEWMAN. 1967. A song sparrow preying on adult Brephos 
infans and Leucobrephos brephoides (Lepidoptera Geometridae) in Michigan. 
Michigan Entomologist 1(7):245-247. 

Dover, C. 1920. The enemies of butterflies. J. Bombay Nat. Hist. Soc. 27:642-643. 

ENDLER, J. A. 1978. A predator’s view of animal color patterns. Evol. Biol. 11:319- 
364. 

ERICKSON, J. M. 1973. Bird predation on Papilio polyxenes F. (Papilionidae). J. Lepid. 
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FRAZER, J. F. D. & M. ROTHSCHILD. 1960. Defense mechanisms in warningly-colored 
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FRYER, J. C. F. 1913. Field observations on the enemies of butterflies in Ceylon. Proc. 
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HEGNAUER, H. 1969. Passafloraceae in chemotaxonomie der Pflanzen V. Bitkhauser 
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JEFFORDS, M. R., J. G. STERNBURG & G. P. WALDBAUER. 1979. Batesian mimicry: Field 
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KNOWLTON, G. F. 1953. Predators of Vanessa cardui. Lepid. News 17(2):55. 

KOLYER, J. M. 1968. Note on damaged specimens. J. Res. Lepid. 7:105-111. 

MarsH, N. & M. ROTHSCHILD. 1974. Aposematic and cryptic Lepidoptera tested on 
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MORLEY, J. & M. SCHACHTER. 1963. Acetylcholine in non-nervous tissues of some 
Lepidoptera. J. Physiol. 168:706. 

Morrison, M. L. 1980. Seasonal aspects of the predatory behavior of loggerhead 
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OLSON, W. L. 1962. Song sparrows feeding on Lepidoptera. J. Lepid. Soc. 16(2):1136. 

PLATT, A. P., R. P. COPPINGER & L. P. BROWER. 1971. Demonstration of the selective 
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PouGcu, F. H. & L. P. BROWER. 1977. Predation by birds on the great southern white 
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27. 

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ROBBINS, R. K. 1978. Behavioral ecology and evolution of hairstreak butterflies (Lep- 
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1980. The lycaenid “false head” hypothesis: Historical review and quantitative 
analysis. J. Lepid. Soc. 34(2):194—208. 


VOLUME 39, NUMBER 4 261 





1981. The “False Head” hypothesis: Predation and wing pattern variation of 
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Pp. 


Journal of the Lepidopterists’ Society 
39(4), 1985, 262-265 


A NEW SPECIES OF CLEARWING MOTH, 
CARMENTA LAURELAE (SESIIDAE), FROM FLORIDA 


Larry N. BROWN 
Department of Biology, University of South Florida, Tampa, Florida 33620 


THOMAS D. EICHLIN 


Insect Taxonomy Laboratory, A. & I., Division of Plant Industry, 
California Department of Food and Agriculture, Sacramento, California 95814 


AND 


J. WENDELL SNOW 
Fruit and Tree Nut Research Laboratory, U.S.D.A., Byron, Georgia 31008 


ABSTRACT. A new species of clearwing moth, Carmenta laurelae, was discovered 
in west-central Florida using the synthetic pheromone, (Z,Z) 3, 18-octadecadien-1-ol 
acetate and is herein described. The type series consists of 75 male specimens which 
were taken only in cypress swamps and the adjacent forested floodplain habitats. The 
flight period of the adult males occurred from 1100-1400 h daily. Adults were captured 
only between 13 May to 3 June 1985 in the Tampa Bay area. 


Although the North American sesiids have been the subject of two 
monographs (Beutenmuller, 1901; and Engelhardt, 1946), the fauna is 
still imperfectly known. The development of several chemical sex at- 
tractants by Tumlinson and colleagues (1974) and Schwarz and col- 
leagues (1983) has greatly aided the collecting of clearwing species and 
also helped elucidate their ecological and taxonomic relationships 
(Duckworth & Ejichlin, 1977). 

While using a variety of sesiid sex attractants to survey for the moths 
in the Tampa Bay area of west-central Florida during 1985, a new 
species of clearwing moth was captured in May and early June. The 
description of this species follows. 


Genus Carmenta Hy. Edwards 
Carmenta laurelae, new species 


Male (Fig. 1): Head. Front brown-black, white laterally; vertex brown-black; occipital 
fringe yellow laterally, orange-yellow and brown-black mixed dorsally; labial palps 
smoothly scaled, dark brown dorsally and laterally, orange-yellow ventrally; antennae 
brown-black, some pale yellow powdering dorso-medially. 

Thorax. Brown-black with narrow subdorsal orange-yellow stripes; orange-yellow patch 
beneath wings. Legs brown-black, with much orange-yellow on forecoxa, near tibial 
spurs, on first tarsal segments and at tarsal joints. 

Wings. Forewing slightly more than one-half hyaline but with broad brown-black 
apical region, leaving small circular hyaline area just distad of discal spot, discal cell and 
region below Cu hyaline, no light scaled powdering dorsally; ventrally with costal margin 
orange-yellow and some orange-yellow powdering between veins on apical region. 
Hindwing hyaline; ventrally with some orange-yellow on costal margin. Forewing length: 
9-10 mm. 


VOLUME 39, NUMBER 4 263 





: 


Fic. 1. Carmenta laurelae, adult male. 


Abdomen. Mostly brown-black, dorsally with narrow orange-yellow band at posterior 
edge of segment 2, broader but still narrow band on posterior margin of segment 4, very 
narrow band on posterior edge of segment 7; ventrally with narrow orange-yellow bands 
on posterior margin of segments 4—7; anal tuft somewhat wedge-shaped but truncate at 
apex, with some orange-yellow at tips of lateral scales and ventrally on tip of abdomen. 

Male genitalia. As illustrated (Fig. 2), with saccus somewhat uncharacteristic for species 
of Carmenta, being only about one-third total length of valve and bilobed apically but 
with other features typical for the genus. 

Female. Unknown. 

Host plant. Unknown. 

Distribution. Florida. 

Types. Holotype: Male, University of South Florida Ecological Research Area, Tampa, 
Hillsborough County, Florida, V-13-1985, Coll. L. N. Brown, ZZ-3, 18 ODDA phero- 
mone; deposited in National Museum of Natural History (NMNH), Washington, D.C. 
Paratypes: 74 males, deposited in NMNH; California Department of Food and Agricul- 
ture (CDFA), Sacramento; Florida State Collection of Arthropods, Gainesville; and au- 
thor’s collection, University of South Florida, Tampa. 


Discussion. This species is only known from the 75 male specimens 
of the type series, which were taken at five different localities in Hills- 
borough County, Florida. The first specimen appeared in a sticky trap 
on 13 May 1985, and the last individual was taken 3 June 1985 (a flight 


264 


LW 
Nie 
AW 


i 
a 
WII 

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mA 
Mp 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Carmenta laurelae, male genitalia (ventral view, left valve removed). 


period of three weeks). All individuals were captured within cypress 
swamps and adjacent floodplain forests. Carmenta laurelae was never 
taken at any pheromone isomer other than (Z,Z) 3, 13-ODDA, although 
several other sex attractants were constantly available. 

Pheromone traps were checked at hourly intervals for several days 
to determine the duration of the daily flight period of male C. lauralae. 


VOLUME 39, NUMBER 4 265 


They came to traps only between 1100-1400 h, with the greatest flight 
activity occurring around noon. 

This moth is named in honor of Laurel Brown, only daughter of the 
collector of the new clearwing species. 


ACKNOWLEDGMENTS 


We thank Charles S. Papp, Sierra Graphics and Typography, Sacramento, California, 
for final inking of the drawing and for photographing the adult moth. Also, thanks go 
to Isa Montenegro, Agricultural Biological Technician, CDFA, Sacramento, for various 
technical assistance. 


LITERATURE CITED 


BEUTENMULLER, W. 1901. Monograph of the Sesiidae of North America, north of 
Mexico. Mem. Am. Mus. Nat. Hist. 1:217-352. 

DuckwortTH, W. D. & T. D. EICHLIN. 1977. Two new species of clearwing moths 
(Lepidoptera: Sesiidae) from eastern North America clarified by sex pheromone. J. 
Lepid. Soc. 31:191-196. 

ENGELHARDT, G. P. 1946. The North American clear-wing moths of the family Ae- 
geriidae. U.S. Natl. Mus. Bull. 190:1-222. 

SCHWARZ, M., J. A. KLUN, B. A. LEONHARDT & D. T. JOHNSON. 1983. (E,Z)-2, 13- 
Octadecadien-1-ol acetate. A new pheromone structure for sesiid moths. Tetrahedron 
Letters 24:1007-1010. 

TUMLINSON, J. H., C. E. YONCE, R. E. DOOLITTLE, R. R. HEATH, C. R. GENTRY & E. 
R. MITCHELL. 1974. Sex pheromones and reproductive isolation of the lesser peach- 
tree borer and peachtree borer. Science 185:614-616. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 266-267 


THE HOST PLANT, ERYTHROXYLUM (ERYTHROXYLACEAE), 
OF AGRIAS (NYMPHALIDAE) 


THOMAS S. RAY 


School of Life & Health Sciences, University of Delaware, 
Newark, Delaware 19716 


ABSTRACT. A male of Agrias amydon philatelica DeVries was reared from its host 
plant, Erythroxylum fimbriatum Peyritsch, in the wet Caribbean lowland forests of 
Heredia Province, Costa Rica. This is the first host plant record for Agrias amydon. 


A male of Agrias amydon philatelica DeVries (DeVries, 1980) was 
reared from its host plant Erythroxylum fimbriatum Peyritsch, at Fin- 
ca La Selva, an Organization for Tropical Studies field station in the 
wet Caribbean lowland forests of Heredia Province, Costa Rica. This 
is the first host plant record for Agrias amydon. 

In March 1979, a penultimate instar larva was found feeding on a 
2.5 m individual of E. fimbriatum in the forest. The plant was a 
member of a clump of five individuals of E. fumbriatum within a space 
of 100 m along the Holdridge Trail. The plants have been vouchered 
and deposited in the Duke University Herbarium, Hammel 8929, Kress 
76-526. The larva was fed in the lab on leaves from the same plant. 
Pupation occurred on the upper surface of the cage. The adult emerged 
in April. The last larval shed skin and head capsule, the empty chrys- 
alis, and the preserved adult were deposited in the collections of the 
Museum of Comparative Zoology of Harvard University. The speci- 
men was designated a paratype by DeVries (1980). 

Previous records of the subspecies, all since 1977, were from the 
Pacific lowland dry forests in Guanacaste Province, Costa Rica, the 
Caribbean lowland wet forests of Herrera Province, Panama, and Fin- 
ca La Selva (DeVries, 1980). This suggests that the butterfly has a wide 
geographical distribution and occurs in widely varying habitats, though 
it must be very rare. 

Subsequent to my observations, D. Janzen and W. Hallawachs (pers. 
comm.) raised A. amydon philatelica from Erythroxylum havanense 
Jacq., at Parque Santa Rosa in Guanacaste. In Guanacaste, E. hava- 
nense is one of the most common shrubs; thus, the rarity of Agrias in 
Guanacaste cannot be explained in light of the abundance of its host 
plant. However, the recent discovery of Agrias in Guanacaste marks 
the first record of Agrias outside of wet forests. 

In view of the long history of collecting in Costa Rica, it is surprising 
that Agrias amydon was first collected there in 1977, although there 
have been at least nine subsequent records (DeVries, 1980) from Cen- 
tral America. This suggests that there may have been a recent increase 


VOLUME 39, NUMBER 4 267 





Fic. 1. Developmental stages of Agrias amydon philatelica (left to right): adult (top), 
larva (bottom), adult emerging from chrysalis, and chrysalis. 


in abundance, and/or possibly a range extension. We know that Agrias 
is capable of using two species of Erythroxylum as its host plant. It 
may also be possible that Agrias is capable of using the commercial 
species Erythroxylum coca Lam. and E. novogranatense (Morris) Hi- 
eron. as its host plants. This is very speculative, and it must be consid- 
ered that the commercial species have higher alkaloid contents. How- 
ever, it has been noted that Agrias is very common in the Tingo Maria 
region of Peru (C. Pringle, pers. comm.), known as a hot spot for the 
production of cocaine. Agrias sardanapalus claudina “‘claudianus’’ is 
reported to feed on Quiiana glaziovi (Barselou, 1983). 


LITERATURE CITED 


BARSELOU, PAUL E. 1983. The genus Agrias. Sciences Nat., Compiegne. 125 pp. 

DEVRIES, P. 1978. A record of Agrias amydon (Nymphalidae) from Costa Rica. J. 
Lepid. Soc. 32(4):310. 

1980. The genus Agrias (Lepidoptera: Nymphalidae: Charaxinae) in Costa 

Rica. Brenesia 17:295-302. 





Journal of the Lepidopterists’ Society 
39(4), 1985, 268-275 


EGG DISPERSION PATTERNS AND EGG AVOIDANCE 
BEHAVIOR IN THE BUTTERFLY 
PIERIS SISYMBRII BDV. (PIERIDAE) 


TIMOTHY A. KELLOGG 


San Francisco State University, San Francisco, California 94132 


ABSTRACT. Egg dispersion patterns of the pierid butterfly, Pieris sisymbrii Bdv., 
were studied in the Mojave Desert, San Bernardino County, California. A census of eggs 
found on its cruciferous hostplant, Arabis pulchra, revealed a clumped egg dispersion 
pattern. This is an unexpected result if females of P. sisymbrii avoid conspecific eggs. 
An egg removal experiment suggested that P. sisymbrii females avoided plants bearing 
eggs, selecting egg-free hosts instead. In times of limited egg-laying sites, extended female 
flights due to successive rejections of egg-bearing plants may be a cue to inhibit egg- 
avoidance behavior, causing females to select egg-bearing hosts more frequently. 


Ovipositing females of some inflorescence or infructescence-feeding 
pierid butterflies avoid those cruciferous host plants bearing conspecific 
eggs. Females which can recognize the presence of conspecific eggs on 
potential host plants apparently avoid host overload by assessing their 
egg-load and if necessary, adjusting their oviposition behavior, accept- 
ing only plants without eggs (Shapiro, 1980). When sufficient host plants 
are available, egg-load assessment and egg avoidance behavior should 
lead to a uniform egg dispersion pattern. Supporting studies have been 
obtained from laboratory work with Pieris brassicae L. (Rothschild & 
Schoonhoven, 1977), and field work with Anthocaris sara Lucas (Sha- 
piro, 1980), Euchloe hyantis Edw. (Shapiro, 198la), and Anthocaris 
cardamines L. (Wiklund & Ahrberg, 1978). 

Prevention of host overload is adaptive in that the amount of food 
plant on a single plant or stem is usually enough for only the first 
hatched larva, and any subsequent larvae may have little or no food 
plant available to consume (Rausher, 1979). Older larvae may also 
exhibit cannibalistic tendencies toward eggs and younger larvae (Stamp, 
1980). When plants free of eggs are available, it would seem maladap- 
tive for assessing butterflies to utilize plants already bearing eggs. 

Several investigators, however, have found clumped or aggregated 
egg dispersion patterns, usually as a consequence of female butterflies 
utilizing isolated plants or plants found along the margins of host plant 
clumps (Mackay & Singer, 1982; Cromartie, 1975; Jones, 1977; Court- 
ney & Courtney, 1982). The use of a few isolated plants by females 
can leave individual hosts with far more eggs than they can support. 

The California white, Pieris sisymbrii Bdv., is a member of a large 
group of pierid butterflies that feed preferentially on inflorescences; 
however, the favorite oviposition sites are on stems and undersides of 
cauline leaves. In the Mojave Desert, P. sisymbrii lays blue-green eggs 


VOLUME 39, NUMBER 4 269 


on its host plant, Arabis pulchra Jones (Brassicaceae). The eggs then 
turn a conspicuous bright orange within a day. Many assessing pierid 
butterflies have brightly colored orange-to-red non-cryptic eggs, per- 
haps to facilitate egg recognition by females (Shapiro, 1981a). 

This study examines egg-load assessment and egg avoidance behav- 
ior in P. sisymbrii in the Mojave Desert. I attempt to answer two 
questions: (1) Is the initial egg dispersion pattern in the field the ex- 
pected uniform distribution, and if not, are there explanations for de- 
viations from uniformity? (2) Does P. sisymbrii avoid egg-bearing host 
plants when egg-free plants are available, thus suggesting a discrimi- 
natory behavior? 


Study Area 


The study site consisted of low rolling hills approximately 1000 m 
in elevation and located 30 km south of Baker, San Bernardino County, 
California (Mojave Desert). Dominant shrubs include Joshua tree (Yuc- 
ca brevifolia Engelm.) and creosote bush (Larrea divaricata Cav.) and 
their associates. The area supports large populations of crucifers, in- 
cluding Caulanthus cooperii (Wats.) Pays., Descurainia pinnata Walt., 
as well as A. pulchra. Arabis pulchra occurs primarily along washes 
between adjacent hills. The plant is perennial and may be woody or 
herbaceous, depending on its age and growing conditions. Its habit is 
variable; occasionally a single plant may have up to 15 stems while 
other plants may have only a single stem. 

Pieris sisymbrii adults are found in open, exposed areas where solar 
radiation is high and are most active in the mid-morning (Emmel & 
Emmel, 1978). 


METHODS 


A sample of 60 specimens of A. pulchra was selected for an egg- 
removal experiment. Plants were chosen based on the following cri- 
teria: (1) High relative conspicuousness; some plants occurred in the 
center of shrubs, making their inflorescences and flowers difficult to 
see. These plants were excluded. (2) Plants with only a minimal amount 
of herbivore damage. No plants with chewed-up leaves or flowers were 
selected. (3) Satisfactory number of flowers and flower buds. Females 
may utilize only those plants with a sufficient food resource for the 
larvae. 

All sampled plants were inspected initially for P. sisymbrii eggs, and 
the number of eggs per plant was recorded. Egg color and egg position 
on the plant were also tabulated. Plants were assigned to a control 
group and an experimental group. Plants bearing at least one egg were 
tagged and assigned to the control group, with their eggs left in place. 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Distribution of Pieris sisymbrii eggs on Arabis pulchra on a per-plant basis. 


All eggs Orange eggs only 
No. eggs/plant No. of plants No. eggs/plant No. of plants 

0 23 0 40 
] 22 al 15 
7 8 2 2 
3 2 3 2 
4 3 4 in 
5 I 60 
6 0 
7 “i 

60 


Plants without eggs were tagged and assigned to the experimental 
group. No apparent morphological differences between egg-free and 
egg-bearing plants were observed. Thirty-seven plants were initially 
found to bear eggs, and to simplify the statistics involved, eggs were 
removed from seven randomly selected control plants, and these plants 
were added to the experimental group. In this way, each group in- 
cluded 30 plants. Eggs were removed from the plants by teasing with 
an insect pin and a camel hair brush. 

The study site was visited for six consecutive days, 27 March through 
1 April 1983. Inspection for newly laid eggs started at 1000 h each day 
just as butterfly activity was peaking. If new eggs were found on the 
control plants they were noted and left on the plant; if new eggs were 
found on the experimentals, they were noted and removed from the 
plant, as described above. A variance-to-mean ratio (s?/x) was used to 
measure dispersion patterns. 


RESULTS 


Sixty-eight eggs were found initially on 37 sample plants, with 15 
of these plants bearing more than one egg. Distribution of eggs on a 
per plant basis appears on Table 1. One individual plant (with a single 
stem) bore seven eggs, all of which were a few inches from one another. 

A variance-to-mean ratio was calculated on a per-plant basis and 
was significantly greater than one, indicating an aggregated egg dis- 
tribution (Table 2). This suggests that P. sisymbrii was not assessing its 
egg-load on A. pulchra. If egg color is an indication of the age of the 
egg, females may be avoiding orange eggs only since these would be 
the first to hatch, and the resulting larvae would have a considerable 
head start in development. The variance-to-mean ratio for orange eggs 
only, however, still deviates toward an aggregated distribution, albeit 
not as extreme a deviation as on a per-plant basis (Table 2). 


VOLUME 39, NUMBER 4 271 


TABLE 2. Dispersion of Pieris sisymbrii Bdv. eggs found on Arabis pulchra Jones. 
Variance-to-ratios (s?/x) were calculated on a per-plant and per-inflorescence basis. 


N x ? (s?/x)* x? 
Per-plant 
All-eggs 60 1.13 1.98 1.75 103.25: 
Orange eggs only 60 0.55 0.73 1.33 78.50? 
Per-inflorescence 
All-eggs 190 0.36 0.68 1.89 359.108 
Orange eggs only 190 0.15 0.24 1.60 302.40 
~*Any value <1 indicates a uniform distribution; >1 indicates an aggregated distribution, ~ indicates a random 
distribution. 
10.005 > P; df = 


20.10 > P > 0.05: ee 
30.005 > P; df = 189 


Shapiro (1980) suggests that assessment works on a per-inflorescence 
basis, where multiple ovipositions occur on plants with many stems. 
Females may perceive individual stems as individual oviposition sites 
and flowers from single inflorescences as adequate food resource for 
one developing larva. In such cases, many eggs may be found on multi- 
stemmed plants. A total of 190 stems were counted from the 60 sample 
plants (Table 3). The deviation toward an aggregated distribution, 
however, was the most extreme in this treatment (Table 2). Over 50% 
of the eggs occurred on stems with at least one other egg. 

The result of the egg removal experiment indicates that P. sisymbrii 
prefers to oviposit on host plants without eggs. During the week, 27 
new eggs were found on the experimental plants compared with 10 on 
the controls (x? = 7.81, df = 1, 0.010 > P > 0.005). This suggests that 
females can recognize eggs and discriminate against those plants bear- 
ing eggs, thus effectively avoiding possible larval competition and in- 
creasing larval survivorship. 


DISCUSSION 


In all treatments, P. sisymbrii eggs in the field failed to correspond 
to the uniform dispersion pattern expected if females show egg-load 
assessment and egg avoidance behavior. The variance-to-mean ratios 
based on a per-plant, egg color, and inflorescence basis all deviated 
toward aggregated egg distributions. Such concentration of eggs on a 
few host plants has been shown to result in lower larval survival in the 
papilionid Battus philenor feeding on Aristolochia (Rausher, 1979). 
The use of a single stem or plant by many larvae will affect their 
chances of survival, especially if the amount of foodplant available to 
an individaul offspring is small relative to its food requirements, or if 
the needed resource is limited by certain age and physiological con- 


AY JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 8. Distribution of Pieris sisymbrii eggs on Arabis pulchra on a per-inflores- 
cence basis. 


All eggs Orange eggs only 

0 146 0 169 
i 28 ] 15 
2 12 2 4 
3 3 3 nil 
4 0 190 
5 0 
6 0 
7 Eh 

190 


ditions, as inflorescences are. Any positive preference for plants bearing 
conspecific eggs would cause intense intraspecific competition and low- 
ered larval survivorship. Such behavior would be maladaptive and re- 
sult in a reduced fitness and therefore should not evolve. 

Several authors have found that individual host plants along the 
margins of clumps may receive a disproportionate number of eggs 
relative to inner plants (Courtney & Courtney, 1980; Mackay & Singer, 
1982; Cromartie, 1975; Jones, 1977). Shapiro (1975) suggests that the 
use of these margin plants would be a selected response to defend 
against parasitoids which key in on high host plant density. Courtney 
and Courtney (1982) studied aggregated egg patterns of Anthocaris 
cardamines but did not prove any single factor as causing dispropor- 
tionate egg concentrations. Several aspects of female behavior, they 
believe, contribute to the large egg loads on individual margin plants— 
or the ‘edge-effect’ as they call it. Such behaviors include (1) a tendency 
of A. cardamines to oviposit on its host plant after flying a long dis- 
tance, even if conspecific eggs are already present, and (2) a host 
searching behavior in which females sample plant clumps as if plant 
density is low or plants are widely dispersed. These behaviors may be 
adaptive if host plant populations undergo periodic fluctuations in total 
biomass. The selective disadvantage of laying eggs on plants with con- 
specific eggs would be counterbalanced by successful discovery of hosts 
during times of low plant density, with females accepting even those 
which have conspecific eggs. 

At the Mojave site, A. pulchra occurred almost exclusively along dry 
washes where plants were distributed in a linear fashion making it 
difficult to visualize a margin. Interplant distance was approximately 
20 m, implying no true area with a high concentration of plants. This 
high interplant distance, however, may cause females to perceive in- 


VOLUME 39, NUMBER 4 2S 


dividual hosts as isolated plants, and large egg loads may be expected. 
Shapiro (198la) found isolated host plants of Barbarea verna (Mill.) 
Asch. to bear up to 15 eggs of Anthocaris sara. Mackey and Singer 
(1982) found that the probability of Euptychia libye L. (Satyridae) 
Ovipositing on a sprig of Panicum sp. increased as the spatial isolation 
of the plant increased. They interpreted their results as being a con- 
sequence of random initiation of search pattern by the female after an 
oviposition period, rather than an active preference for isolated plants. 
It was difficult to judge if any particular plant at the Mojave site was 
more isolated than others. All plants, however, were within sight of 
each other, and no plants were found outside the wash. 

Despite the initial aggregated egg dispersion pattern, P. sisymbrii 
preferred to oviposit on host plants free of eggs, according to the results 
of the egg removal experiment. The experimental plants received 27 
new ovipositions within the week compared with 10 on the controls. 
One experimental plant received a new oviposition on three consecu- 
tive days. Two other experimental plants, on the second and fifth day 
respectively, bore two new green eggs during the day; both were count- 
ed as a single oviposition event. 

The aggregated egg dispersion patterns and the apparent egg avoid- 
ance behavior seem to offer conflicting evidence. Yet, aggregated dis- 
persion patterns need not disprove some degree of egg avoidance be- 
havior (Singer & Mandracchia, 1982). My data is consistent with the 
selective behavior proposed by Courtney and Courtney (1982) where 
long flight distances by females could cause them to lose their discrim- 
inating behavior and lay more eggs on normally avoided plants. When 
suitable host plants are sparse due to adverse environmental conditions, 
such as a flood or drought, females have abnormally long oviposition 
flights to find the few plants available and ignore any eggs that are 
present. Therefore, indiscriminate egg-laying behavior after lengthy 
oviposition flights may be advantageous in habitats with great environ- 
mental extremes where the probability of encountering egg-free plants 
is sometimes low. The consequence of encountering and rejecting egg- 
laden plants consecutively is an increase in flight time and flight dis- 
tance. This could be the cue which serves to inhibit assessment behavior 
and increase the probability of a female accepting an oviposition site 
that would normally be disregarded. 

Possible coevolutionary results of egg-load assessment and egg-avoid- 
ance behavior can be seen on a local race of one of P. sisymbrii’s 
northern California host plants, Streptanthus brewerii Gray (Shapiro, 
1981b). These plants produce pigmented callosites which appear to be 
“egg-mimics.’’ Females were shown to be more apt to oviposit on plants 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


which had their callosites removed, thus, suggesting a visual cue for 
egg avoidance (see also Williams & Gilbert, 1981). 

My study assumes that assessment behavior is primarily based on 
visual cues, although pheromonal ovipositional deterrents have been 
observed in a butterfly species (Rothschild & Schoonhoven, 1977) and 
in a few dipteran species (Prokopy, 1975; Zimmerman, 1979) and 
therefore cannot be ruled out. 

Other possible responses exist besides the inhibition of avoidance 
behavior. In times of limited optimal oviposition sites, females may 
disperse to another suitable area or choose alternative hosts. Further 
study is needed to check for any relationships between extended ovi- 
positional flights and the degree of host selectivity by the females. It 
is suggested, however, that P. sisymbrii may avoid those plants bearing 
conspecific eggs as long as egg-free plants are available as alternatives. 
Once all or most of the available hosts have been taken, females may 
cease their discriminatory behavior and oviposit on plants with eggs 
instead of not laying eggs at all. 


ACKNOWLEDGMENTS 


This study was part of a graduate independent research project at San Francisco State 
University. I wish to thank the CSU Consortium Field Laboratory at Zzyzx Springs, 
California for use of their facilities; my advisor, Dr. John Hafernik, Jr., for his guidance; 
Leslie Watson for her assistance in the field and on the typewriter; Albert Wilson for his 
encouragement and sense of humor; and Dr. Arthur Shapiro of the University of Cali- 
fornia at Davis for his expertise and valuable suggestions. 


LITERATURE CITED 


COURTNEY, S. P. & S. COURTNEY. 1982. The edge effect in butterfly oviposition: cau- 
sality in Anthocaris cardamines and related species. Ecol. Entomol. 7:131-1387. 
CROMARTIE, W. J. 1975. The effect of stand size and vegetational background on the 

colonization of cruciferous plants by herbivorous insects. J. Appl. Ecol. 12:517-538. 
EMMEL, T. C. & J. F. EMMEL. 1973. The butterflies of southern California. Natural 
History Museum of Los Angeles County Publications, Los Angeles, California. 
JONES, R. E. 1977. Movement patterns and egg distributions in cabbage butterflies. J. 

Anim. Ecol. 46:195-212. 
Mackay, D. A. & M. C. SINGER. 1982. The basis of an apparent preference for isolated 
host plants by ovipositing Euptychia libye butterflies. Ecol. Entomol. 7:299-303. 
Proxkopy, R. J. 1975. Ovipositing-deterring fruit marking pheromone in Rhagoletis 
fausta. Environ. Entomol. 4:298-300. 

RAUSHER, M. D. 1979. Egg recognition: its advantage to a butterfly. Anim. Behav. 27: 
1034-1040. 

ROTHSCHILD, M. & L. M. SCHOONHOVEN. 1977. Assessment of egg load by Pieris 
brassicae (Lepidoptera: Pieridae). Nature 266:352-355. 

SHAPIRO, A. M. 1975. Ecological and behavioural aspects of coexistence in six crucifer- 
feeding Pierid butterflies in the central Sierra Nevada. Am. Midl. Nat. 93:424—4383. 

1980. Egg-load assessment and carry over diapause in Anthocaris (Pieridae). 

J. Lepid. Soc. 4:307-315. 

198la. The Pierid red-egg syndrome. Am. Nat. 117:276-294. 








VOLUME 39, NUMBER 4 OTD 


1981b. Egg-mimics of Streptanthus (Cruciferae) deter oviposition by Pieris 
sisymbrii (Lepidoptera: Pieridae). Oecologia 48:142-143. 

SINGER, M. C. & J. MANDRACCHIA. 1982. On the failure of two butterfly species to 
respond to the presence of conspecific eggs prior to oviposition. Ecol. Entomol. 7: 
327-330. 

STAMP, N. E. 1980. Egg deposition patterns in butterflies: Why do some species cluster 
their eggs rather than deposit them singly? Am. Nat. 115:367-380. 

WIKLUND, C. & C. AHRBERG. 1978. Host plants, nectar source plants, and habitat 
selection of males and females of Anthocaris cardamines (Lepidoptera). Oikos 31: 
169-183. 

WILLIAMS, K. S. & L. E. GILBERT. 1981. Insects as selective agents on plant vegetative 
morphology: egg mimicry reduces egg laying by butterflies. Science 212:467—469. 

ZIMMERMAN, M. 1979. Oviposition behavior and the existence of an ovipositing-deter- 
ring pheromone in Hylemya. Environ. Entomol. 8:277-279. 





Journal of the Lepidopterists’ Society 
39(4), 1985, 276-279 


THE FOOD PLANTS OF JALMENUS DAEMELI SEMPER 
(LYCAENIDAE) WITH NOTES ON OTHER BUTTERFLIES 
AND ACACIA FOOD PLANTS 


T. J. HAWKESWOOD 


49 Venner Road, Annerley, Brisbane, 
Queensland 4108, Australia 


ABSTRACT. The literature providing larval food plant data for Jalmenus daemeli 
Semper (Lycaenidae) is summarized. A new larval host plant, Acacia leucoclada Tindale 
subsp. argentifolia Tindale (Mimosaceae) is recorded from the Warwick district, south- 
eastern Queensland. One previously overlooked host, A. pendula A. Cunn. ex G. Don, is 
included here from the published literature. The name Acacia cunninghamii Hook. is 
no longer valid as the food plant for four butterflies, Jalmenus evagoras (Donovan), J. 
daemeli Semper, J. ictinus Hewitson and Hypochrysops delicia delicia Hewitson, since 
the revised classification of Acacia in Queensland does not allow accurate determinations 
for the food plants referred to under the name cunninghamii. Comments are made on 
a new host recorded for J. evagoras. The known larval hosts for J. daemeli are 12 and 
for J. evagoras 15. 


Jalmenus daemeli Semper (Damel’s blue) occurs from Cairns to 
Brisbane in scattered localities along the coast and also in certain inland 
localities such as Eidsvold, Gayndah, Toowoomba, Stanthorpe and Mil- 
merran (Common & Waterhouse, 1972, 1981). Atkins (1976) recorded 
J. daemeli from various localities in central Queensland, while De Baar 
(1977) recorded it from an area between Bunya Mountains and Ar- 
chookoora State Forest in southeastern Queensland. The species is note- 
worthy in usually having large, isolated populations. Little has been 
published on its biology. Following the convention of an earlier paper 
on the larval food plants of Jalmenus evagoras (Donovan) (Hawkes- 
wood, 1981), the known larval hosts of J. daemeli are listed and dis- 
cussed below. 


Larval Host Plants 


The first records of Acacia (Mimosaceae) being listed as larval food 
plants appears to be those of Lucas (1889) and Illidge (1898). They 
noted that J. daemeli (Ialmenus illidgei Lucas, in the case of Lucas, 
1889) fed on wattles in the Brisbane area, southeastern Queensland. 
Gurney (1911) also stated the species fed on wattles. However, none 
of these authors provided specific determinations for these plants. I]- 
lidge (1921) recorded myall (Acacia pendula A. Cunn. ex G. Don) as 
a larval food plant from the Jandowae district, southeastern Queens- 
land. This record was overlooked by Common and Waterhouse (1972, 
1981). Illidge (1921) noted that the butterfly was abundant in all stages 
on young myall trees and were attended by ants. Manski (1960) re- 
corded Acacia cunninghamii Hook. as a food plant from Marybor- 


VOLUME 39, NUMBER 4 277 


ough, Scarborough and Redcliffe (the latter two localities are now outer 
suburbs of Brisbane). Waterhouse (1932:190) recorded brigalow (Aca- 
cia harpophylla F. Muell. ex Benth.) as a host, while Common (1964: 
92) recorded A. harpophylla and “other wattles” and Heterodendrum 
(Sapindaceae). Macqueen (1965) recorded J. daemeli as occasionally 
attacking Heterodendrum diversifolium F. Muell. (Sapindaceae). He 
also noted that in the Toowoomba district, southeastern Queensland, 
J. daemeli fed solely on the silver-leaf ironbark, Eucalyptus melano- 
phloia F. Muell. (Myrtaceae) and another unidentified species of blood- 
wood, Eucalyptus sp., despite Acacia being plentiful in the district. 
Harslett (1965) recorded Acacia neriifolia A. Cunn. ex Benth., A. de- 
currens (Wendl.) Willd. and A. irrorata Sieb. ex Spreng. as food plants 
from Stanthorpe, southeastern Queensland. It should be noted that A. 
decurrens (green wattle) is endemic to New South Wales and Victoria 
and is naturalized near Toowoomba and Stanthorpe (Pedley, 1978; 
Stanley & Ross, 1983). Atkins (1975) recorded Acacia bidwillii Benth. 
(erroneously cited as Acacia bidwelli Benth.) as a larval host from four 
localities in central Queensland, viz. Rockhampton, Thompson’s Point, 
Wycarbah and Broadsound Range. He also recorded A. bancroftii 
Maiden and A. macradenia Benth. as food plants from the Expedition 
Range, central Queensland. Lane (1979) noted, “It is of interest that 
Acacia bidwillii has also been observed as a food plant of J. daemeli 
Semper in numerous localities between Rockhampton and Mackay, 
Queensland,” but he did not provide a reference. Presumably his com- 
ments are based on observations by Atkins (1975). I have also observed 
larvae and pupae on the leaves and stems of young A. bidwillii plants 
(about 1 m high) growing on the James Cook University campus, 
Townsville, north Queensland, during 16-26 November 1981. They 
were associated with large numbers of an Iridomyrmex (Hymenoptera: 
Formicidae). Adult butterflies visited the open flowers of A. bidwillii 
(Hawkeswood, 1985). Mr. M. De Baar (June 1984, pers. comm.) has 
recorded large numbers of larvae and pupae of J. daemeli on Acacia 
leucoclada Tindale subsp. argentifolia Tindale, 25 km south of War- 
wick, southeastern Queensland during January 1988. This is a previ- 
ously unrecorded host for this butterfly. 

Manski (1960) recorded A. cunninghamii Hook. as a larval host for 
four species of Lycaenidae—Hypochrysops delicia delicia Hewitson, 
Jalmenus evagoras evagoras (Donovan), J. ictinus Hewitson and J. 
daemeli Semper (noted above). (Waterhouse (1932) originally recorded 
this Acacia for H. d. delicia). However, in a recent revision of the 
Queensland Acacia species, Pedley (1978) noted that the name A. cun- 
ninghamii had been applied loosely to six Acacia species, viz. A. tro- 
pica (Maiden et Blakely) Tindale, A. cretata Pedley, A. longispicata 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Benth. (ssp. longispicata and velutina Pedley), A. crassa Pedley (ssp. 
crassa and longicoma Pedley), A. concurrens Pedley and A. leiocalyx 
(Domin) Pedley (ssp. leiocalyx and herveyensis Pedley). (As a result 
of Pedley’s revision, the name cunninghamii should not be used for 
any Acacia). Therefore, the records of A. cunninghamii as a host could 
apply to any of the above six species. In respect to J. daemeli, J. 
evagoras and J. ictinus, the observations by Manski (1960) were made 
in the Maryborough and Brisbane districts, while those of H. d. delicia 
were restricted to Maryborough. Of the six Acacia species noted above, 
only two, A. crassa longicoma and A. leiocalyx leiocalyx, are known 
to occur naturally in the Maryborough district, while A. concurrens 
and A. leiocalyx leiocalyx grow in the Brisbane district (from Pedley, 
1978). Since A. leiocalyx is usually more common in both districts, it 
is possible that the name A. cunninghamii, referred to in Manski (1960), 
refers to this species. Whether J. daemeli and the other butterflies 
utilize A. concurrens, A. leiocalyx and A. crassus, or a combination of 
these, must await the results of further field work. In the meantime, 
the name A. cunninghamii listed in Waterhouse (1932), Manski (1960), 
Common and Waterhouse (1972, 1981) and Hawkeswood (1981) should 
be disregarded. 

For J. daemeli, there are 12 known larval host plants, viz. Acacia 
bancroftii, bidwillii, decurrens, harpophylla, irrorata, leucoclada subsp. 
argentifolia, macradenia, neriifolia, pendula, Heterodendrum diver- 
sifolium, Eucalyptus melanophloia and Eucalyptus sp. 

In reference to Jalmenus evagoras, Dunn (1984) recently recorded 
a new larval host, Acacia diffusa Ker (erroneously cited as Acacia 
diffusa Lindl.). This species is regarded as a synonym of Acacia genis- 
tifolia Link (Jacobs & Pickard, 1981). With A. cunninghamii omitted 
from my list (i.e., Hawkeswood, 1981) and A. genistifolia included, 
the number of larval hosts remains at 15, all of which are Acacia 
species. 


ACKNOWLEDGMENTS 


I would like to thank Mr. M. De Baar, Department of Forestry, Indooroopilly, Bris- 
bane, Queensland, for his unpublished information on J. daemeli, for discussions on the 
general biology of butterflies and their food plants and for pointing out several references. 
I also thank Mr. A. Hiller, Mt. Glorious, Queensland for information on J. daemeli. This 
research was undertaken on private funds. 


LITERATURE CITED 


ATKINS, A. F. 1975. Larval food plants of some Queensland butterflies. News Bull. 
Entomol. Soc. Qd. 3:117-119. 

1976. New records for butterflies in southern, central and northern Queensland. 
Aust. Entomol. Mag. 3:1-4. 

Common, I. F. B. 1964. Australian butterflies. Jacaranda Press, Brisbane. 131 pp. 





VOLUME 39, NUMBER 4 279 


Common, I. F. B. & D. F. WATERHOUSE. 1972, 1981. Butterflies of Australia. Angus 
& Robertson, Sydney. Ist ed. 498 pp. (1972), 2nd Ed. 682 pp. (1981). 

DE Baar, M. 1977. Butterflies from an area between the Bunya Mountains and Ar- 
chookoora State Forest, Queensland. Aust. Entomol. Mag. 3:115-119. 

Dunn, K. L. 1984. Acacia diffusa Lind].—A new larval foodplant for Jalmenus eva- 
goras evagoras (Donovan) (Lepidoptera: Lycaenidae). Vict. Entomol. 14:8. 

GURNEY, W. B. 1911. A study of wattle trees (Acacia) and a list of insects of wattle 
trees. Aust. Nat. 2:56-59. 

HARSLETT, J. 1965. Butterflies from the Stanthorpe district, Queensland, with notes on 
their food plants. Qd. Nat. 17:106-112. 

HAWKESWOOD, T. J. 1981. The food plants of Jalmenus evagoras (Donovan) (Lepi- 
doptera: Lycaenidae). Aust. Entomol. Mag. 8:1-2. 

1985. The role of butterflies as pollinators of Acacia bidwillii Benth. (Mimo- 
saceae) at Townsville, north Queensland. Aust. J. Bot. 33:167-173. 

ILLIDGE, R. 1898. List of butterflies of the Brisbane district. Proc. Roy. Soc. Qd. 13: 
89-102. 

1921. Rhopalocera of the Jandowae district of the Darling Downs. Qd. Nat. 3: 
23-24. (See also Errata, Qd. Nat. 3:48.) 

Jacoss, S. W. L. & J. PICKARD. 1981. Plants of New South Wales. Government Printer, 
Sydney. 226 pp. 

LANE, D. A. 1979. Life history notes and distribution records for some Queensland 
butterflies. Aust. Entomol. Mag. 5:115-117. 

Lucas, T. P. 1889. Six new species of Rhopalocera. Proc. Roy. Soc. Qd. 6:155-161. 

MACQUEEN, J. 1965. Notes on Australian Lycaenidae (Lepidoptera). J. Entomol. Soc. 
Qd. 4:56-57. 

MANSKI, M. J. 1960. Food plants of some Queensland Lepidoptera. Qd. Nat. 16:68-73. 

PEDLEY, L. 1978. A revision of Acacia Mill. in Queensland. Austrobaileya 1:75-234. 

STANLEY, T. D. & E. M. Ross. 1983. Flora of south-eastern Queensland. Vol. I. Queens- 
land Dept. of Primary Industries, Misc. Publ. 81020, Government Printer, Brisbane. 
545 pp. 

WATERHOUSE, G. A. 1932. What butterfly is that? Angus & Robertson, Sydney. 291 


PP. 








Journal of the Lepidopterists’ Society 
89(4), 1985, 280-283 


PREDATION ON CATOCALA MOTHS (NOCTUIDAE) 


AUBURN E.. BROWER 
8 Hospital St., Augusta, Maine 04330 


ABSTRACT. Catocala adults are preyed upon by flying squirrels; tree and fence 
lizards; bats and birds, commonly in man-made situations. 


I was born in the last century, in a family with natural history 
interests on the Ozark Mountain uplift in southwest Missouri. My first 
interest was ornithology/oology, then collecting insect and plant ma- 
terials for sale, exchange and my collections. From the back porch I 
could look across a small field at the edge of 320 acres of “wild land,” 
never cut, never fenced. The upland tree community was largely post, 
black and blackjack oak; the hollows had hickory, white and red oak 
with other trees especially along streams. The woods were commonly 
burned each spring, it was said, to kill the ticks, snakes and to rejuve- 
nate the grass. The burning resulted in many of the trees having basal 
fire scars. On poor soil with a deficiency of rain, the timber was small, 
open and more widely spaced with limited underbrush. This presented 
an ideal situation in which to observe animal and insect life. Oats were 
sown in March and, like winter wheat, cut and shocked in June. Then, 
there was much time for me to be afield on my interests, in near 
continuously warm sunny days. That was Catocala country every year. 
Without electricity, light and baits yielded few specimens, and so, 
daylight collecting was carried on. 

I have over 200 predator-injured Catocala, including every one of 
the larger eastern species. Injuries to their wings are various. I think 
Dr. T. Sargent did a good job of illustrating in his book “Legion of 
Night, The Underwing Moths” a theoretical injury which could be 
produced by the attack of a bird. But, in all of my injured Catocala, 
there is not one injury I think was produced by a bird peck. I once 
saw a flycatcher fly near a fluttering Catocala while popping its beak, 
probably because the bird’s nest was near. The Catocala I have found 
resting on trees in depressions in the bark on oak trees draw their wings 
down tight, and I have said that a bird would need a vanadium steel 
lower mandible to bite a piece out of the wing or wings on one side. 
In the big woods of northwest Maine I once saw a scarlet tanager up 
in the top of a tall white birch, and the wings of a Catocala fluttered 
down. The tanager may have devoured the moth. A. J. Snyder says 
that he saw a Catocala “‘snapped” out of a tree by a scarlet tanager 
and immediately torn to pieces. Of utmost importance, fully 70% of 
all specimens of some lots injured have similar injuries in all four wings 


VOLUME 39, NUMBER 4 281 


or in the two hindwings, most of these obviously made at the same 
moment by the predator. These types of injury were not made by 
birds, even to flying moths. 

Flying squirrels, genus Glaucomys, seem ideally fitted to attack Ca- 
tocala as they live in the same situations. They are the most active 
voracious predators I know. One year I prepared a stupifying bait, fed 
through wicks, with a screen wire basket below to catch the moths 
which fell. All I could get were wings, and all I found accountable 
were flying squirrels, which I knew to be strongly insectivorous. I took 
jump-steel traps, attached noctuid moths to the pan and hung them 
on the tree trunks above the bait outlet. In these traps I caught flying 
squirrels, one after another, until I gave up that collecting method. 
They are widespread along with Catocala, where old trees remain in 
some numbers. The above mentioned injuries to the Catocala spp. are 
common. 

In the Ozarks I have watched skinks or tree lizards flushing out the 
moths up on the limby portion of tree trunks, where Catocala tend to 
rest in the morning and on overcast days. Wings were found on the 
ground in such areas. These lizards are well fitted to attack moths and 
produce in an instant the torn pairs of wings which are so prevalent 
where the lizards live. Florida collectors have spoken of these. A con- 
siderable number of moths which Dean Berry sent me from Florida 
had over 40% with serious injury to their wings. 

Conditions are different in other parts of the country. John W. John- 
son of Corona del Mar, California, a shrewd experienced worker, writes: 
“...in fifty years of field collecting Catocala I have seen little evidence 
of predation. I have never witnessed an attack by birds on a flying 
Catocala, nor observed them searching tree boles for the moths. Nor 
have I ever collected moths showing the types of wing damage figured 
in Sargent’s “Legion of Night” that has been ascribed to birds. An 
experienced observer in the 1920’s and 1930’s, Janet Riddell, saw a 
lizard, Sceloporus sp. hunting over a tree hole, encountering a Catocala 
at rest which it seized by the body, bit off the wings close to the body 
and swallowed the body whole. I have frequently observed Sceloporus 
sp. lizards climbing about tree boles as high as 15-20 feet above the 
ground, and have found sets of Catocala wings clipped off close to the 
body at the base of trees in groves where Catocalas were present and 
resting, which I supposed due to lizard predation.’’ He has repeatedly 
mentioned the fact that the smaller western Catocala regularly fly 
from one thick clumped scrub oak to another two feet or less above 
ground, thus avoiding most predators. He collected a Catocala cali- 
forniensis Brower with a spine of a low cactus driven 4 mm into its 
thorax. 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Dave Baggett has kindly forwarded an extensive account of numer- 
ous experiences with predation on Catocala in Florida. His experiences 
usually include man’s changes and their effects on both predatory life 
and Catocala, which differ greatly from my experiences while in my 
teens and twenties in the Ozark mountain area. He says: “... while 
eating lunch several C. ilia Cramer wings filtered down from the tree 
tops near me, and another specimen from which the abdomen had 
been chewed off, but the wings and thorax were still intact—the moth 
still quivering. The birds eating them were blue jays, but we never saw 
one of these birds catch one.” He says that in Jacksonville his bait trap 
area was a regular feeding area for mockingbirds, English sparrows 
and blue jays. As his traps were emptied of insects early each morning, 
the birds would rapidly snap up the majority of the moths as they flew 
away. The birds preferred smaller moths as a rule; preferring Catocala 
amica Htibner and micronympha Gueneé over ultronia Hubner, ilia, 
muliercula Gueneé, etc., all common species. He says he has seen many 
wild specimens with bird beak patterns. He suspects the heaviest pre- 
dation is by lizards, especially anoles and fence swifts. Even though 
they are relatively small, they will readily grab at large Catocala like 
ilia, lacrymosa Gueneé, agrippina Strecker, etc., frequently getting 
completely yanked off the tree trunk by the frantic flapping of the 
moth to get away, and most of the larger ones probably do. One rarely 
finds the smaller species with the rounded “‘lizard-type’’ bites on the 
wings. He has repeatedly seen American anoles capture and eat Ca- 
tocala moths, including species as large as ultronia; and the anoles try 
to catch larger species, frequently leaving their marks. While trying 
to collect C. jair Strecker, an eastern fence swift on an oak was ob- 
served catching and eating one of these moths which, when flushed 
out, had settled near the lizard. Flying squirrels in Florida most defi- 
nitely catch and eat Catocala and sometimes also gray squirrels. Some 
who have used bait traps or have baited trees will confirm this. Dave 
Baggett says on numerous occasions while watching lighted sheets he 
has seen bats and owls capture larger moths and Catocala spp., also at 
city lights, always catching the moths in mid-air. He has seen red- 
shouldered hawks catch the larger saturniid moths. He thinks birds do 
not aggressively seek out Catocala selectively. Certainly, that is not the 
case in Florida. Presumably, the greater number of mangled Catocala 
in collections have been attacked by lizards and not by the more pow- 
erful flying squirrels which can destroy a much larger number of the 
insects. At Ithaca, New York, skunks visited both the Cornell light trap 
and baited Crataegus shrubs. 

Flying squirrels and tree and fence lizards are the important pred- 
ators on Catocala. Moths at light and coming to bait attract skunks and 


VOLUME 39, NUMBER 4 283 


other animals. Bats are important predators around man-lighted areas. 
Instances of bird predation are reported for blue jays, scarlet tanagers 
and man-flushed Catocala by wood pewee. Many common species of 
birds are attracted to the insects which are flushed from light, bait and 
artificial traps. Birds sieze numerous small insects when traps are cleared 
releasing the insects, attracted as to bird feeders. Hornets and wasps 
also take part. 


LITERATURE CITED 


SARGENT, THEODORE D. 1976. Legion of night, the underwing moths. University of 
Massachusetts Press, Amherst. 222 pp. 

SNYDER, A. J. 1897. A remarkable appearance of Catocala insolabilis Gueneé. Can. 
Entomol. 29:70. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 284-298 


AN ANNOTATED LIST OF THE BUTTERFLIES AND 
SKIPPERS OF LAWRENCE COUNTY, OHIO 


JOHN V. CALHOUN 
6332 C Ambleside Dr., Columbus, Ohio 43229 


ABSTRACT. Until recently, only 23 species of butterflies and skippers were known 
from Lawrence County. In 1983-1984 a study was conducted to increase our knowledge 
of these insects in the county. As a result, 60 additional species were recorded. One 
species, Euchloe olympia, was recorded in Ohio for the first time. For each species listed, 
the following data are provided: relative abundance, habitat and nectar sources, extreme 
dates, and localities. Species recorded prior to this study are accompanied by historic 
collection data. A list of 21 additional species which should be looked for in Lawrence 
County is included. Twenty-four species are not known from the adjacent counties in 
Kentucky or West Virginia. Thirteen species showed differences in abundance between 
1983 and 1984 and potential reasons are discussed. Curves are also provided to illustrate 
relative species diversity during the study. The county possesses characteristics more 
typical of regions to the south of Ohio and the Appalachian uplands, and several resident 
species of butterflies and skippers reflect these aspects. A list of species found in Lawrence 
County, Ohio allows for a more complete understanding of the butterfly and skipper 
fauna of southern Ohio, northeastern Kentucky and southwestern West Virginia. 


Two lists have dealt with the butterflies and skippers of southeastern 
Ohio (Parshall, 1983; Shuey, 1983). Although Vinton and Athens coun- 
ties have received attention, other counties in the region have virtually 
been ignored. The natural history of Lawrence County is poorly known, 
and Albrecht (1982) recorded only 23 species of butterflies and skippers 
from the county. The southernmost county in Ohio, Lawrence County 
is relatively inaccessible and remains insufficiently understood. In 1983 
a study was conducted to increase our knowledge of the butterfly and 
skipper fauna of Lawrence County. As a result, 41 additional species 
were recorded. The study was continued in 1984 and 19 previously 
unrecorded species were collected, bringing the total number of species 
known from the county to 83. One species, Euchloe olympia (W. H. 
Edwards), represents a state record and brings the total number of 
species recorded from Ohio to 137 (Riddlebarger, 1984). The present 
paper describes the results of this study and provides historic collection 
data on the butterflies and skippers of Lawrence County, Ohio. 

Lawrence County is situated on the Ohio River, bordering the states 
of Kentucky and West Virginia (Fig. 1). The average annual temper- 
ature of the county is approximately 138°C (Gordon, 1969). The average 
annual precipitation ranges from 102-112 cm with snowfall measuring 
approximately 38-64 cm (Collins, 1975). Frost dates for spring (dates 
after which there is a 50% or less chance that temperatures will fall to 
O°C or lower) are 20-25 April, and for fall (dates by which there is a 
50% chance that the first O°C temperature will have occurred) are 15- 
20 October (Collins, 1975). Thick river fogs may contribute to a mod- 


VOLUME 39, NUMBER 4 285 













LAKE 
ESUVIUS 





WINDSOR ATHALIA®: 


ROME | 







UNION 

PROCTORVILLE 

CHESAPEAKE = 

FAYETTE aes 

ef, SOUTHS” BEN 
=P 


os 












Fic. 1. The location, political divisions, cities, and collecting localities of Lawrence 
County, Ohio. Land within the boundaries of Wayne National Forest is shaded. 


eration in climate immediately along the Ohio River (Cusick & Sil- 
berhorn, 1977). 

Lawrence County lies within the Unglaciated Allegheny Plateau 
Region of Ohio. Relative relief ranges from 91-152 m (Gordon, 1969). 
Underlain by a bedrock of Pennsylvanian shale and sandstone, the 
original vegetation of the county at the time of the earliest land surveys 
consisted of mixed oak (Quercus spp.) forests with limited tracts of 
mixed mesophytic and bottomland hardwood forests (Fig. 2). Mixed 
oak forests covered knobs and ridgetops and were composed of a chest- 
nut oak (Quercus montana Willd.)—chestnut (Castanea dentata (Marsh) 
Borkh.) forest type (Gordon, 1969). Yellow pines (Pinus rigida Mill., 
Pinus virginiana Mill., and Pinus echinata Mill.) occurred locally 
(Cusick & Silberhorn, 1977). Mixed mesophytic forests greatly varied 
in composition and were dominated mostly by broad-leaved species, 
with no single species comprising a large fraction of the dominants 
(Gordon, 1966). This forest type was found on less well drained north 
and northeast facing slopes (Cusick & Silberhorn, 1977). Bottomland 
hardwood forests were variable in composition and occurred in older 


286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 







MIXED OAK 
cet MIXED MESOPHYTIC 


[_|BOTTOMLAND 
HARDWOOD 





Fic. 2. The original vegetation of Lawrence County (adapted from Gordon, 1966). 


valleys and on recent alluvium and terraces of major streams (Gordon, 
1966). Intense lumbering during the 19th century, developmental pres- 
sures, and agricultural activities have significantly altered and frag- 
mented these forests. Consequently, no pristine woodland communities 
remain. Today, the county is more than 70% forested and composed 
of oak-hickory (Carya sp.), Virginia pine (Pinus virginiana)-pitch 
pine (Pinus rigida), and oak—pine forest types (Fig. 3). 

Dean State Forest and the Ironton District of Wayne National Forest 
presently contain the most extensive tracts of mature secondary wood- 
land in Lawrence County. Here, controlled clear-cutting and strip min- 
ing have created many habitats of various successional stages. The 
federally administered Lake Vesuvius Recreation Area possesses a man- 
made lake surrounded by a forest with a rich floral composition. Else- 
where in the county private woodlots of secondary forest, cropland, 
pastures, and fallow fields are the principal types of vegetational com- 
munities (Fig. 4). Wetlands are few, but several buttonbush, Cepho- 
lanthus occidentalis L., and cat-tail, Typha sp., marshes and thinly 


VOLUME 39, NUMBER 4 287 







-OAK—HICKORY 
[_ JOAK—PINE 


—— =: VIRGINIA—PITCH 
PINE 





Fic. 3. The current major forest types of Lawrence County (adapted from Ohio 
Dept. of Nat. Res., 1984). 


wooded swamps of willow, Salix spp., and river birch, Betula nigra 
L., exist in the northern portion of the county. 

Due to its location, Lawrence County possesses characteristics more 
typical of regions south of Ohio. As a result of a long growing season 
and low winter precipitation, a few species of plants are known in 
Ohio only from bluffs along the Ohio River in Lawrence County and 
the adjacent counties of Gallia and Scioto (Cusick & Silberhorn, 1977). 
One southern species, American holly, Ilex opaca Ait., is believed to 
occur naturally in Ohio only in Lawrence and Scioto counties (Braun, 
1961). Lawrence County also exhibits affinities to the Appalachian up- 
lands. Several species of Rhododendron, limited in Ohio to the more 
rugged part of the Unglaciated Allegheny Plateau, occur locally in the 
county (Braun, 1961). 

The butterfly and skipper fauna of Lawrence County was undoubt- 
edly modified when the original vegetation was destroyed. Prior to 


288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 4. Habitats in Lawrence County. (Top) Ridgetop oak-pine forest (Wayne Na- 
tional Forest). (Bottom) Disturbed area along the Ohio River (South Point). 


VOLUME 39, NUMBER 4 289 


settlement, species adapted to forest habitats probably predominated. 
Presettlement Athens County was also heavily forested and Shuey (1983) 
speculated that species characteristic of open areas possibly were absent 
or inhabited ephemeral areas. A comparable situation could have ex- 
isted in Lawrence County. Today, the county has a more diverse flora 
and thus, probably supports a more diverse butterfly and skipper fauna 
as well. 


METHODS 


The results are based upon 30 visits to Lawrence County during 19 
July-7 October 1983 and 27 April—11 October 1984. Historic records 
were gathered from Stehr (1945), Albrecht (1982; pers. comm., 1983), 
and the 1979 Lepidopterists’ Society Season Summary (News of the 
Lepid. Soc., No. 2. Mar/Apr, 1980). Although collecting was done 
throughout the county, Wayne National Forest and areas along the 
Ohio River received the most attention. 

The results are presented in the following format: species name, 
relative abundance, habitat and nectar sources, extreme dates, and 
localities. Designations of relative abundance are adapted from Covell 
(1984). These designations apply only when weather conditions are 
favorable and individuals are most active. “Abundant”? means that a 
species can be expected in great numbers in the correct habitat and 
season. “Common” indicates that a species can be expected in the 
correct habitat and season, and several specimens can be anticipated. 
“Uncommon” means that a species may or may not be found in the 
proper habitat and season; or that very few specimens might be found 
on a given visit to a specific location. ““Rare” species are seldom en- 
countered. Distinct differences in the abundance of a species between 
1983 and 1984 are shown by listing two abundance designations, one 
for each year, separated by a slash symbol. Habitat information is based 
upon observations made by the author in Lawrence County. Nectar 
source information is provided for species seen visiting flowers and 
helps to indicate correct habitat associations. Participation in mud pud- 
dling behavior is also noted. Extreme dates are given to show the 
approximate length of flight periods during the study but are not con- 
clusive. Dates are organized as day/month/year. Only positive iden- 
tifications of specimens captured or observed in the field were used in 
compiling extreme dates. Specific localities are given for species usually 
encountered singly or that exist in very localized colonies. Localities 
refer to those on Fig. 1. Species recorded from Lawrence County prior 
to the 19838-1984 study are accompanied by historic collection data in 
parentheses. The taxonomy and nomenclature follow A Catalogue / 
Checklist of the Butterflies of America North of Mexico (Miller and 


290 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brown, 1981), and each species listed is preceded by the number used 
in that publication. An asterisk (*) denotes a species that Albrecht 
(1982) and Opler (1983) do not record from the adjacent counties in 
Ohio (Gallia, Jackson, and Scioto), Kentucky (Boyd and Greenup), or 
West Virginia (Cabell and Wayne). However, one of these species, 
Staphylus hayhurstii (W. H. Edwards), was discovered by the author 
in Jackson County, Ohio in 1984. Specimens collected by the author 
are contained in his private collection and the Ohio Historical Society 
collection in Columbus, Ohio. 


RESULTS 


HESPERIOIDEA 
HESPERIIDAE 


7a Epargyreus c. clarus (Cramer). Abundant. Hayfields, forest clearings, roadsides; 


40 


42 


47 
48 


AHO 


83 
84a 
85a 


90 


92 
AS 


115 


131 


red clover (Trifolium pratense L.), common milkweed (Asclepias syriaca L.); 
mud puddles. 11.vi.84—8.x.83. Throughout. (1944, Hanging Rock, A. W. Lind- 
sey). 

Autochton cellus (Boisduval and LeConte). Uncommon. Ridgetop trail through 
oak forest, forest margins, roadsides; New Jersey tea (Ceanothus americanus L.), 
wild ipicac (Gillenia stipulacea (Muhl. ex Willd.)), purple milkweed (Asclepias 
purpurascens L.), common milkweed; mud puddles. 19.vi.84—2. vii.84. Blackfork, 
Decatur Twp., Lake Vesuvius. (16.vi.71, D. K. Parshall). 

Achalarus lyciades (Geyer). Uncommon. Dry fields, brushy roadsides, margins 
of oak forests; red clover, common milkweed. 11.vi.84—30.viii.83. Throughout. 
(29.v.1899, collector unknown). 

Thorybes bathyllus (J. E. Smith), Uncommon/common. Disturbed areas near oak 
forest; red clover. 11.vi.84—29.viii.83. Throughout. (26.vi.32, J. S. Thomas). 
Thorybes pylades (Scudder). Common. Disturbed areas near woods; red clover. 
11.vi.84-17.vii.84. Throughout. (17.vi.1899, J. S. Thomas). 

Staphylus hayhurstii (W. H. Edwards). Uncommon. Gardens, wooded stream 
banks, and trails near the foodplant, Chenopodium album L.; yellow wood sorrel 
(Oxalis sp.); mud puddles. 29. vii.84—9.viii.83. Sybene, South Point, Coal Grove. 
Erynnis icelus (Scudder and Burgess). Rare. Margins of oak forest; mud puddles. 
11.vi.84-19.vi.84. Blackford, Decatur Twp. 

Erynnis b. brizo (Boisduval and LeConte). Rare. Margins of oak forest; mud 
puddles. 25.iv.84-28.iv.84. Washington Twp. 

Erynnis j. juvenalis (Fabricius). Common. Oak forest and margins; mud puddles. 
25.iv.84-5.v.84. Throughout. 

Erynnis horatius (Scudder and Burgess). Common/uncommon. Oak forest and 
margins; wild ipicac; mud puddles. 25.iv.84—29.viii.83. Washington Twp., De- 
catur Twp., Lake Vesuvius. 

Erynnis martialis (Scudder). Rare. One female in a dry ridgetop clearing near 
the foodplant, Ceanothus americanus. 12.vi.84. Decatur Twp. 

Erynnis baptisiae (Forbes). Rare. Disturbed areas along the Ohio River; ironweed 
(Vernonia sp.), red clover, 9.viii.83—8.x.83. Proctorville, South Point. 

Pholisora catullus (Fabricius). Common. Hayfields and gardens near the food- 
plant, Chenopodium album, alfalfa (Medicago sativa L.), dogbane (Apocynum 
sp.), red clover. 26.vi.84—4.ix.84. Throughout. (1.vi.1899, Collector unknown). 
Nastra lherminier (Latreille). Rare. Brushy field and roadside where the food- 
plant, Andropogon scoparius Michx., is common. 11.vi.84—28.viii.83. Washington 
Twp. 


VOLUME 39, NUMBER 4 291 


*142 
*151 


161 


174 
179 
180a 


185 


*186 


187b 


197 


198 


*217b 


*219a 
*945 


*259 


297a 


300 


308a 
*314 


320a 


325a 


Ancyloxypha numitor (Fabricius). Common. Wet fields, wooded clearings near 
streams; red clover. 11.vi.84-17.ix.84. Throughout. 

Hylephila phyleus (Drury). Rare. Disturbed areas along the Ohio River; white 
Aster sp., alfalfa. 4.viii.84—8.x.83. Proctorville, South Point, Ironton. 

Hesperia leonardus Harris. Rare/Locally common. Ridgetop old fields, clearings, 
pastures; thistle (Cirsium sp.), ironweed. 14.viii.83—12.ix.84. Blackfork, Washing- 
ton Twp., Decatur Twp. 

Polites coras (Cramer). Abundant. Any open area; red clover, alfalfa, ironweed; 
mud puddles. 11.v.84—8.x.83. Throughout. 

Polites themistocles (Latreille). Uncommon. Hayfields, roadsides; red clover. 
11.v.84-28. viii.83. Throughout. 

Polites o. origenes (Fabricius). Uncommon/common. Hayfields, dry old fields, 
forest clearings; red clover. 11.vi.84—12.ix.84. Throughout. 

Wallengrenia egeremet (Scudder). Rare. Oak ridgetop clearings, margin of oak 
forest; mud puddles. 25.vi.84—5.viii.84. Decatur Twp., Lake Vesuvius. (30. vi.34, 
J. S. Thomas). 

Pompeius verna (W. H. Edwards). Common. Fallow fields, clearings, brushy 
roadsides; red clover, common milkweed. 19.vi.84—29.vii.84. Throughout. 
Atalopedes campestris huron (W. H. Edwards). Abundant/common. Any open 
area; red clover, ironweed, asters; mud puddles. 17.vii.84—11.x.84. Throughout 
in 1988, only along Ohio River in 1984. (2.viii.82, Union Twp., J. V. Calhoun). 
Poanes hobomok (Harris). Rare. Grassy clearings, oak forest margins. 11.vi.84- 
26.vi.84. Blackfork, Lake Vesuvius. 

Poanes zabulon (Boisduval and LeConte). Uncommon. Grassy clearings, oak 
forest margins. 26.vii.83-5.viii.84. Blackfork, Lake Vesuvius, Ironton. (21.v.1899, 
A. W. Lindsey). 

Euphyes ruricola metacomet (Harris). Uncommon. Hayfields, old fields, road- 
sides; red clover, common milkweed; mud puddles. 11.vi.84—-17.viii.84. Through- 
out. 

Atrytonopsis h. hianna (Scudder). Rare. Ridgetop oak forest clearings where the 
foodplant Andropogon scoparius, occurs. 11.vi.84—12.vi.84. Decatur Twp. 
Amblyscirtes vialis (W. H. Edwards). Rare. Trail through ridgetop oak forest; 
mud puddles. 5.v.84. Lake Vesuvius. 

Panoquina ocola (W. H. Edwards). Rare. One specimen in disturbed area along 
Ohio River; white Aster sp. 7.x.83. South Point. 


PAPILIONOIDEA 
PAPILIONIDAE 


Battus p. philenor (Linnaeus). Uncommon/common. Forests, forest margins, 
hayfields, roadsides; joe-pye-weed (Eupatorium fistulosum Barratt), red clover, 
common milkweed; mud puddles. 25.iv.84-17.ix.84. Throughout. (26.vi.32, C. F. 
Walker). 

Eurytides marcellus (Cramer). Rare. Forests, forest margins; purple loosestrife 
(Lythrum salicaria L.), wild ipicac. 25.iv.84—2.viii.83. Washington Twp., Lake 
Vesuvius, Coal Grove. (27.vi.79, F. Bower). 

Papilio polyxenes asterius Stoll. Uncommon. Hayfields, pastures; red clover. 
11.vi.84—12.ix.84. Throughout. 

Heraclides cresphontes (Cramer). Rare. Forest margins, hayfield; red clover, 
common milkweed. 4.viii.84—14.viii.83. Lake Vesuvius, Proctorville. 

Pterourus g. glaucus (Linnaeus). Abundant. Forests, forest margins, hayfields, 
roadsides; dandelion (Taraxacum officianale Weber), red clover, thistle, common 
milkweed, joe-pye-weed; mud puddles. 25.iv.84—12.ix.84. Throughout. (27.vi.79, 
F. Bower). 

Pterourus t. troilus (Linnaeus). Abundant. Forests, forest margins, hayfields, 
roadsides; red clover, thistle, common milkweed, joe-pye-weed; mud puddles. 
25.iv.84—12.ix.84. Throughout. 


292 


*334 


338 
*344 


349b 


dolla 


352 


383a 


*388 


39la 


393a 
*398 


417b 


424b 


*427b 


44] 


468a 


470a 


478b 


503a 


505a 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PIERIDAE 


Pontia protodice (Boiduval and LeConte). Uncommon. Disturbed areas along 
the Ohio River; red clover, dogbane, ironweed. 17.vii.84—11.x.84. Athalia, South 
Point. 

Artogeia rapae (Linnaeus). Common. Nearly any open area; red clover, alfalfa, 
asters; mud puddles. 25.iv.84—11.ix.84. Throughout. 

Euchloe olympia (W. H. Edwards). Rare. Oak forest ridgetop. 28.iv.84—-5.v.84. 
Lake Vesuvius. 

Falcapica midea annickae dos Passos and Klots. Uncommon. Forests, forest mar- 
gins, 27.iv.84-5.v.84. Decatur Twp., Lake Vesuvius. (18.iv.76, Washington Twp., 
C. W. Albrecht). 

Colias p. philodice Godart. Common. Hayfields, vacant lots, roadsides; red clover, 
alfalfa, asters; mud puddles. 25.iv.84—-11.x.84. Throughout. (18.iv.76, Washington 
Twp., C. W. Albrecht). 

Colias eurytheme Boisduval and LeConte. Common. Hayfields, vacant lots, road- 
sides; red clover, alfalfa, asters; mud puddles. 5.v.84-11.x.84. Throughout. 
Pyrisitia |. lisa (Boisduval and LeConte). Common/rare. Clearings, dry fields, 
roadsides; mud puddles. 19.vii.83-11.x.84. Throughout in 1983, only South Point 
in 1984. 

Abaeis nicippe (Cramer). Rare. Brushy field near the foodplant, Cassia hebecarpa 
Fern., trail through ridgetop oak forest, grassy roadside. 5.v.84—12.ix.84. Wash- 
ington Twp., Decatur Twp., Lake Vesuvius. 


LYCAENIDAE 


Feniseca t. tarquinius (Fabricius). Uncommon. Clearings and sunlit stream banks 
and lanes near common alder (Alnus serrulata (Ait.) Willd.) and hawthorns 
(Crataegus sp.) infested with wooly aphids (Eriosomatidae); mud puddles. 
19.vi.84—5.viii.84. Decatur Twp., Lake Vesuvius, South Point. 

Lycaena phlaeas americana Harris. Rare. One specimen in disturbed area along 
Ohio River. 8.x.83. South Point. 

Hyllolycaena hyllus (Cramer). Rare. Wet fields and marshes. 11.vi.84—4.ix.83. 
Symmes Twp., Decatur Twp. 

Harkenclenus titus mopsus (Hubner). Uncommon. Oak forest clearings, clearing 
on wooded stream floodplain; common milkweed. 25.vi.84—2.viii.84. Blackfork, 
Lake Vesuvius. 

Satyrium calanus falacer (Godart). Uncommon. Oak forest clearings and mar- 
gins; common milkweed. 19.vi.84—2.vii.84. Blackfork, Decatur Twp., Lake Ve- 
suvius. (26.vi.32, J. S. Thomas). 

Satyrium liparops strigosum (Harris). Rare. Oak forest clearing and margin. 
19.vi.84-25.vi.84. Blackfork, Lake Vesuvius. 

Calycopis cecrops (Fabricius). Uncommon. Oak forest clearings and disturbed 
areas near the foodplant, Rhus copallina L. 11.vi.84-12.ix.84. Blackfork, Lake 
Vesuvius, South Point. 

Incisalia h. henrici (Grote and Robinson). Common. Oak forest clearings and 
margins where the foodplant, redbud (Cercis canadensis L.) occurs; redbud; mud 
puddles. 25.iv.84—-5.v.84. Blackfork, Decatur Twp., Lake Vesuvius. 

Incisalia n. niphon (Hubner). Uncommon. Ridgetops near the foodplant, Pinus 
virginiana; mud puddles. 25.iv.84—29.iv.84. Decatur Twp. 

Strymon melinus humuli (Harris). Rare. Oak forest margins, hayfields; red clo- 
ver, common milkweed. 26.vi.84—4.ix.84. Decatur Twp., Coal Grove. (16.vi.71, 
D. K. Parshall). 

Everes c. comyntas (Godart). Abundant. Hayfields, pastures, roadsides; red clo- 
ver. 5.v.84-11.x.84. Throughout. (18.iv.76, Washington Twp., C. W. Albrecht). 
Celastrina |. ladon (Cramer). Rare/common. Forests, forest margins, clearings; 
redbud; mud puddles. 25.iv.84—4.ix.84. Throughout. (16.vi.71, D. K. Parshall). 


VOLUME 39, NUMBER 4 293 


506 


514d 


*529 


*552a 


*562 


565a 


566a 
580a 


606a 


623b 


635a 
636 
637 
*648a 
*650 
*651 
653a 


*656 


663c 


*664a 


Celastrina ebenina Clench. Uncommon. Margins of lowland forest; mud puddles. 
25.iv.84—5.v.84. Lake Vesuvius. 

Glaucopsyche I. lygdamus (Doubleday). Common. Oak forest margins, clearings, 
and roadsides near the foodplant, wood vetch (Vicia caroliniana Walt.); wood 
vetch; mud puddles. 25.iv.84—5.v.84. Blackfork, Decatur Twp., Lake Vesuvius. 


RIODINIDAE 


Calephelis borealis (Grote and Robinson). Locally abundant. Lowland oak forest 
margins along south-facing roadbank where the foodplant, Senecio obovatus 
Muhl. is common; black-eyed susan (Rudbechia hirta L.), butterfly-weed (Ascle- 
pias tuberosa L.). 25.vi.84-17.vii.84. Lake Vesuvius. 


LIBYTHEIDAE 


Libytheana b. bachmanii (Kirtland). Uncommon/rare. Margins of cropland, sun- 
lit streambanks and wooded lanes near the foodplant, Celtis occidentalis L.; 
tickseed-sunflower (Bidens sp.), asters; mud puddles. 26.vii.83-11.x.84. Lake Ve- 
suvius, South Point, Coal Grove, Hamilton Twp. 


NYMPHALIDAE 


Euptoieta claudia (Cramer). Uncommon. Hayfields, brushy fields; red clover, 
ironweed, asters; mud puddles. 12.vi.84—11.x.84. Symmes Twp., Sybene, South 
Point. 

Speyeria c. cybele (Fabricius). Uncommon/abundant. Hayfields, old fields, forest 
clearings, pastures, roadsides; red clover, common milkweed, ironweed, thistle. 
11.vi.84—11.x.84. Throughout. (27.vi.79, F. Bower). 

Speyeria a. aphrodite (Fabricius). Rare. Oak forest clearings, pastures; common 
milkweed, ironweed. 19.vi.84—4.ix.83. Blackfork, Decatur Twp. 

Clossiana b. bellona (Fabricius). Common. Wet hayfields, vacant lots; red clover. 
17.vii.84-11.x.84. Throughout. (27.vi.79. F. Bower). 

Charidryas n. nycteis (Doubleday and Hewitson). Common. Forest margins, 
sunlit streams and lanes; common milkweed; mud puddles. 11.vi.84—25.viii.83. 
Throughout. (16.vi.71, D. K. Parshall). 

Phyciodes t. tharos (Drury). Abundant. Hayfields, brushy fields, forest clearings, 
roadsides; red clover, alfalfa, common milkweed, butterfly-weed, asters; mud 
puddles. 11.vi.84-11.x.84. Throughout. 

Euphydryas p. phaeton (Drury). Locally common. Thinly wooded swamp, edges 
of cat-tail marsh; mud puddles. 11.vi.84—25.vi.84. Blackfork, Washington Twp. 
Polygonia interrogationis (Fabricius). Uncommon. Oak forests and their mar- 
gins; mud puddles. 19.vi.84—11.x.84. Throughout. 

Polygonia comma (Harris). Common. Oak forests and their margins; mud pud- 
dles. 25.iv.84-11.x.84. Throughout. 

Nymphalis a. antiopa (Linnaeus). Rare. Margins of ridgetop oak forests. 27.vi.84. 
Decatur Twp. 

Vanessa virginiensis (Drury). Uncommon. Hayfields, old fields, roadsides; red 
clover, ironweed, thistle, asters. 25.iv.84—11.x.84. Throughout. 

Vanessa cardui (Linnaeus). Uncommon/absent. Hayfields, roadsides; red clover, 
alfalfa. 19.vii.83—4.ix.83. Throughout. 

Vanessa atalanta rubria (Fruhstorfer). Uncommon. Hayfields, oak forest mar- 
gins, old fields, roadsides; mud puddles. 25.iv.84—5.viii.84. Throughout. 

Junonia coenia Hubner. Common. Hayfields, vacant lots; red clover, ironweed, 
asters; mud puddles. 17.vii.84—-11.x.84. Throughout in 1983, only along Ohio 
River in 1984. 

Basilarchia arthemis astyanax (Fabricius). Common. Forest margins, sunlit lanes; 
mud puddles. 11.vi.84—4.ix.83. Throughout. 

Basilarchia a. archippus (Cramer). Uncommon. Wet fields and marshes near the 
foodplant, Salix sp.; mud puddles. 11.vi.84—17.ix.84. Washington Twp., Symmes 
Twp., Decatur Twp., South Point. (16.vi.71, D. K. Parshall). 


294 


698 


704 


WL 


*718a 


720 


723a 


732c 


760 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


APATURIDAE 


Asterocampa celtis (Boisduval and LeConte). Uncommon. Forest margins and 
lanes near the foodplant, Celtis occidentalis; mud puddles. 11.vi.84—29.viii.83. 
Symmes Twp., South Point, Elizabeth Twp. (1930's or early ’40’s, W. C. Stehr). 
Asterocampa clyton (Boisduval and LeConte). Rare. Forest margins and lanes 
near the foodplant, Celtis occidentalis; mud puddles. 19.vi.84—29.viii.83. Decatur 
Twp., Lawrence Twp., South Point. 


SATYRIDAE 


Enodia anthedon A. H. Clark. Common. Lowland forests and margins, shaded 
grassy swamps; mud puddles. 25.vi.84—29. viii.83. Decatur Twp., Lake Vesuvius, 
Coal Grove. (16.vi.71, D. K. Parshall). 

Cyllopsis g. gemma (Hubner). Rare/uncommon. Ridgetop oak forests, shaded 
grassy swamps; mud puddles. 5.v.84—29.viii.83. Lawrence Twp., Lake Vesuvius. 
(30.vi.34, Dean State Forest, J. S. Thomas). 

Hermeuptychia sosybius (Fabricius). Common. Lowland forest clearings, wood- 
ed stream banks, shaded grassy swamps; mud puddles. 11.vi.84—30.viii.83. 
Throughout. 

Megisto c. cymela (Cramer). Uncommon/common. Forest margins and clear- 
ings. 11.vi.84—5.viii.84. Throughout. (30.vi.34, J. S. Thomas). 

Cercyonis pegala alope (Fabricius). Uncommon. Hayfields, old fields, vacant lots. 
25.vi.84—17.ix.84. Throughout. 


DANAIDAE 


Danaus plexippus (Linnaeus). Common. Hayfields, pastures, vacant lots; red clo- 
ver, common milkweed, ironweed, asters. 25.vi.84—11.x.84. Throughout. 


HYPOTHETICAL SUPPLEMENTARY LIST 


The following list suggests species that should be looked for in Law- 
rence County. Required habitats and foodplants are available in the 
county and these species may occur as breeding residents or strays 
from other regions. An asterisk (*) denotes species that Albrecht (1982) 
and Opler (1983) record from adjacent counties in Ohio, Kentucky, 
and/or West Virginia. 


96 
100b 
104 
150 
165a 
189a 

ZOD 


337 
368a 
3871b 
389 


423 
*425 


HESPERIIDAE 


Erynnis lucilius (Scudder and Burgess). 
Pyrgus centaureae wyandot (W. H. Edwards). 
Pyrgus communis (Grote). 

Thymelicus lineola (Ochsenheimer). 

Hesperia m. metea Scudder. 

Atrytone I. logan (W. H. Edwards). 
Amblyscirtes hegon (Scudder). 


PIERIDAE 


Artogeia virginiensis (W. H. Edwards). 
Zerene c. cesonia (Stoll). 

Phoebis sennae eubule (Linnaeus). 
Nathalis iole Boisduval. 


LYCAENIDAE 


Satyrium edwardsi (Grote and Robinson). 
Satyrium caryaevorum (McDunnough). 


VOLUME 39, NUMBER 4 295 


*460a Mitoura g. gryneus (Hubner). 

464c Incisalia augustus croesoides Scudder. 

*474a Euristrymon o. ontario (W. H. Edwards). 

477 ~Parrhasius m-album (Boisduval and LeConte). 
*491  Erora laeta (W. H. Edwards). 


NYMPHALIDAE 
567  Speyeria idalia (Drury). 
*645  Polygonia progne (Cramer). 
APATURIDAE 
695 Anaea andria Scudder. 
DISCUSSION 


Thirteen species exhibited distinct differences in abundance between 
1983 and 1984. Two species (Hesperia leonardus and Polites origenes) 
showed an increase in 1984 that may be attributed to a lack of col- 
lecting in the proper habitats in 1983. Conversely, another species 
(Libytheana bachmanni) showed a decrease in 1984 that may be at- 
tributed to a lack of collecting in the proper habitats in 1984. Four 
multivoltine species (Thorybes bathyllus, Celastrina ladon, Cyllopsis 
gemma, and Megisto cymela) are more common during their first 
brood, which was not observed in 1983 and consequently, showed a 
marked increase in 1984. Three additional species (Atalopedes cam- 
pestris, Pyrisitia lisa, and Vanessa cardui) are migratory and known 
for their sporatic occurrences in Ohio. All showed a drastic decrease 
in 1984, suggesting that 1983 was a peak year for migrants or 1984 
was poor, or both. The remaining three species either showed a con- 
siderable increase (Battus philenor and Speyeria cybele) or a consid- 
erable decrease (Erynnis horatius) for reasons unknown. 

Fig. 5 presents curves constructed to illustrate seasonal patterns of 
relative diversity during the study period. Visits lasting more than one 
day and visits made on consecutive days are treated as single visits. 
Although several habitat types were sampled during each visit, differ- 
ential collecting undoubtedly caused slight variations in the number of 
species recorded. The curves reveal that the fewest number of species 
occurred in the spring and fall with the greatest number occurring in 
mid to late summer. The lowest total was 11 species observed 1-2 
October 1988, and the highest total was 48 species observed 14-15 
August 1983. Although data on 1983 are incomplete, the relative di- 
versities of the two years are very similar from mid-July through mid- 
October. 

Two species are scarce and very local in occurrence in Ohio yet 
were found in atypically high concentrations in Lawrence County. 
Although listed as uncommon, approximately two dozen individuals of 
Autochton cellus were observed in the county during 1984. This species 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


oO 
i) 


it 


is 
oi 
ro) 





no. of species/v 
w 
ro) 


St a eee 
O “apr may Jn Jy © aug sept oct 


Fic. 5. Relative diversity during the study. 


is usually encountered singly in Ohio. Previously, only a rich valley in 
Hocking County, Ohio called ““Neotoma”’ has yielded a large number 
of specimens from one locality. In Lawrence County, the presence of 
A. cellus was difficult to predict, but up to 11 individuals were seen at 
each of two locations. Calephelis borealis is widespread in Ohio but is 
usually found in low numbers at any given location. However, at the 
single known Lawrence County colony, nearly 100 individuals were 
observed in a single day. Another species, Hesperia leonardus, is most 
often found singly or in very low numbers in Ohio except in the north- 
western portion of the state where large colonies occur. In Lawrence 
County a colony was discovered which contained at least 50 individuals 
in an area approximately 1 hectare in size. In addition, Pontia proto- 
dice has nearly disappeared from Ohio, and the Lawrence County 
record is only the third record in the state since 1966. The county is 
also the only known Ohio location for Atrytonopsis hianna beyond 
Lucas and Fulton counties in the northwestern corner of the state. 
The southern and Appalachian aspects of Lawrence County are re- 
flected in the presence of several resident species of butterflies and 
skippers. Calycopis cecrops, Hermeuptychia sosybius, and Autochton 
cellus are found near the northern limits of their respective distribu- 
tions. The Lawrence County population of Glaucopsyche lygdamus is 
included in the Appalachian distribution isolate of the species recog- 
nized by Opler and Krizek (1984). Long suspected to occur in Ohio, 
Euchloe olympia is known in the state only from Lawrence County. 
The population of this species in the county should be included in the 


VOLUME 39, NUMBER 4 297 


more southern Appalachian distribution isolate of the species recog- 
nized by Opler and Krizek (1984). Celastrina ebenina is closely asso- 
ciated with rich forested Appalachian slopes in the eastern portion of 
its range (Wagner & Mellichamp, 1978). 

Sixty-one percent of the 137 species of butterflies and skippers re- 
corded in Ohio are represented in Lawrence County. One additional 
species, Celastrina neglectamajor Tutt, has recently been recognized 
as distinct from Celastrina ladon (Opler & Krizek, 1984) and may 
occur in Lawrence County. Specimens resembling this species were 
occasionally observed but not collected during 1984, hence this species 
was not included in the list. It is possible that another species, Speyeria 
diana (Cramer), was present in the county prior to settlement and 
subsequently was extirpated. An old specimen of this species labeled 
“southeastern Ohio” is contained in the collection of the Carnegie 
Museum of Natural History. This species inhabits old-growth hard- 
wood forests and suffered a decline within its range when much of the 
habitat was destroyed due to logging and agriculture (Clark & Clark, 
1951). Today, this species may be expanding (Hammon & McCorkle, 
1983(84)). If once present in Lawrence County, it is undoubtedly no 
longer a breeding resident but could become re-established in the fu- 
ture, especially in the maturing woodlands of Wayne National Forest. 

The butterfly and skipper fauna of northeastern Kentucky and south- 
western West Virginia is still only remotely known. Since similar hab- 
itats exist throughout the region, a list of species known to occur in 
Lawrence County, Ohio facilitates a more thorough understanding of 
the butterflies and skippers, not only of southern Ohio, but also of the 
corresponding portions of Kentucky and West Virginia. 


ACKNOWLEDGMENTS 


I wish to thank Dr. David J. Horn of The Ohio State University for critically reviewing 
the manuscript. I also thank Dr. David C. Iftner, John A. Shuey, Julia M. Cornett, Martin 
Hall, Joe Riddlebarger, and members of the Ohio Lepidopterists for their assistance and 
companionship during several visits to Lawrence County. Lastly, I thank Dr. Carl W. 
Albrecht, Jr., for historic collection data. 


LITERATURE CITED 


ALBRECHT, C. W. 1982. The taxonomy, geography, and seasonal distribution of Rho- 
palocera in Ohio. Ph.D. Dissert., The Ohio State Univ., Columbus. 512 pp. 

BRAUN, L. E. 1961. The woody plants of Ohio. Ohio State University Press. 362 pp. 

CLarK, A. H. & L. F. CLarkx. 1951. The butterflies of Virginia. Smithsonian Misc. 
Coll., Vol. 116, No. 7. 239 pp. 

COLLINs, C. W. 1975. Ohio: An atlas. Amer. Printing and Pub. Inc., Madison, Wiscon- 
son. 310 pp. 

COVELL, C. V., JR. 1984. A field guide to the moths of eastern North America. Hough- 
ton-Mifflin, Boston. 496 pp. 


298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cusick, A. W. & G. M. SILBERHORN. 1977. The vascular plants of unglaciated Ohio. 
Ohio Biol. Surv. Vol. V, New Ser. No. 4. 157 pp. 

GORDON, R. B. 1966 Natural vegetation of Ohio at the time of the earliest land surveys. 
Ohio Biol. Surv. Map. 

1969. The natural vegetation of Ohio in pioneer days. Ohio Biol. Surv. Vol. 
III, New Ser. No. 2. 113 pp. 

HAMMOND, P. C. & D. V. MCCORKLE. 1983(84). The decline and extinction of Speyeria 
populations resulting from human environmental disturbances (Nymphalidae: Ar- 
gynninae). J. Res. Lepid. 22(4):217-224. 

MILLER, L. D. & F. M. Brown. 1981. A catalogue/checklist of the butterflies of 
America north of Mexico. Lepid. Soc. Mem. No. 2. 280 pp. 

OHIO DEPT. OF NATURAL RESOURCES. 1984. The future of Ohio’s forests—Looking 
good. Div. of Forestry. 15 pp. 

OPLER, P. A. 1983. County atlas of eastern United States butterflies (1840-1982). Mim- 
eographed. Dist. by the author. 76 pp. 

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the great plains. Johns Hopkins 
Univ. Press, Maryland. 294 pp. 

PARSHALL, D. K. 1983. A primary check list for the butterfly and skipper populations 
of Zaleski State Forest Vinton County Ohio. Mimeographed. Dist. by the author. 4 





pp. 

RIDDLEBARGER, J. E. 1984. Euchloe olympia, a butterfly new to Ohio. Ohio J. Sci. 84: 
267. 

SHUEY, J. A. 1983. An annotated checklist of the butterflies of Athens County, Ohio. 
Ohio J. Sci. 83(5):262-269. 

STEHR, W. C. 1945. Notes and records of Lepidoptera in Ohio. Ohio J. Sci. 45:18. 

WAGNER, W. H., JR. & T. L. MELLICHAMP. 1978. Foodplant, habitat, and range of 
Celastrina ebenina (Lycaenidae). J. Res. Lepid. 32(1):20-36. 


Journal of the Lepidopterists’ Society 
39(4), 1985, 299-312 


SKIPPERS: POLLINATORS OR NECTAR THIEVES? 


B. ADRIENNE B. VENABLES AND EDWARD M. BARROWS 


Department of Biology, Georgetown University, 
Washington, D.C. 20057 


ABSTRACT. The hypothesis that butterflies as a group are primarily nectar thieves, 
rather than pollinators, of many flowers that they visit was tested by observing skippers 
and quantifying their pollen loads. Two species of skippers, Atalopedes campestris and 
Epargyreus clarus, were studied. 

Adult A. campestris visited 23 flower species and Epargyreus clarus visited 27 flower 
species. Fifty-nine male and female E. clarus carried a mean of 45.1, and 283 male and 
female A. campestris carried a mean of 68.4 pollen grains from eight species of very 
frequently visited flowers. Skippers carried most of the pollen in their facial cavities and 
on their proboscides. At least one skipper of each species carried pollen from each of 
these flowers in its genital cavity, a newly documented pollen-carrying structure for 
butterflies. 

The skippers may have occasionally pollinated their nectar flowers, because they were 
constant to particular species during foraging bouts; they transported pollen; and they 
contacted stigmas with their pollen-bearing proboscides. Nevertheless, the skippers evi- 
dently functioned mainly as nectar thieves. They foraged mostly on asteriads rather than 
other kinds of flowers, primarily probing innermost (male-stage) disk florets, and they 
tended not to contact the outermost (female-stage) florets with their more pollen-laden 
parts. Moreover, they carried pollen loads that were too small to make them significant 
pollinators. Thus, our skipper data do not reject the above hypothesis. 


Many butterfly species visit flowers from which they imbibe nectar 
(Faegri & van der Pijl, 1966; Shields, 1972; Barrows, 1976, 1979; 
Schemske, 1976; Wiklund et al., 1979; Schemske & Horwitz, 1984). 
For example, 197 butterfly species found in eastern North America use 
at least 5.9 + 0.55 SE (1-15) genera of flowers as nectar sources (Opler 
& Krizek, 1984). Butterflies undoubtedly pollinate some flower species 
(Grant & Grant, 1965; Levin, 1972; Levin & Berube, 1972; Barrows, 
1979; Cruden & Hermann-Parker, 1979), and they are definitely nectar 
thieves of others (Spears, 1983; Schemske & Horvitz, 1984). An indi- 
vidual nectar thief is an animal that takes nectar through a natural 
orifice of a flower without pollinating it (Inouye, 1980). Further, if an 
animal species thieves nectar during more than 50% of its visits to a 
particular flower species, the entire animal species could be classified 
as a thief species with regard to this plant species. 

Delpino (1874) suggested that male butterflies are likely cross pol- 
linators of their nectar plants, but decades later Robertson (1924: 100- 
101) stated that butterfly ‘relations to flowers are often that of nectar 
thieves.”” Subsequently, Wiklund et al. (1979) studied the flower visit- 
ing of the pierid Leptidea sinapsis L. in Sweden. From this species 
they generalized that, “Butterflies as a group may have evolved to 
occupy a parasitic mode of life as adults, feeding on the nectar of 
flowers without pollinating them,’ but they did not refer to Delpino’s 


300 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


or Robertson’s assertions. All in all, however, pollination effectiveness 
and efficiency of butterflies is little known (Gilbert & Singer, 1975; 
Kevan & Baker, 1983; Spears, 1983). In an attempt to test further the 
butterflies-as-nectar-thieves hypothesis, we studied foraging behavior 
of two common skippers, Atalopedes campestris (Boisduval) and Epar- 
gyreus clarus (Cramer), in Washington, D.C. The identities and rele- 
vant characteristics of the skippers’ nectar flowers, skipper foraging 
behavior, and the locations and amounts of pollen that skippers carried 
were examined to test the hypothesis. Both skipper species that we 
studied are native to the Washington, D.C., area, where they have 
three broods per season (Clark, 1932). Atalopedes campestris fly in the 
garden from mid-July through September; E. clarus, from mid-June 
to early August. A future paper will discuss whether butterflies, in 
general, are nectar thieves or pollinators. 


MATERIALS AND METHODS 


In our study, we define “foraging bout’ as a skipper’s feeding activ- 
ity on one or more flower species, starting when it was first discovered 
on a flower until it could no longer be followed due to its flying out of 
sight. A “visit” is a skipper’s alighting upon or near a flower, extending 
its proboscis into it for at least 1 sec, and presumably feeding. An 
“infrequently visited flower species (IVFS)” is a flower that we saw 
only one individual skipper visit during only one of the ten 2-week 
observation periods of our study. A “frequently visited flower species 
(F VFS)” is a flower that we saw two to four conspecific skippers visit, 
and a “very frequently visited flower species (VF VFS)” is a flower that 
we saw five to hundreds of skippers visit during two or more of the 
2-week observation periods. A “clear day” is one over 75°F, with no 
rain, and with less than 20% cloud cover. A “facial cavity” is a con- 
cavity into which a skipper’s proboscis coils; a “genital cavity,’ one at 
the end of a skipper’s abdomen, formed in a female by scales surround- 
ing her papilla analis above and lamella antevaginalis below and in a 
male by scales surrounding his uncus above and valvae below. 

Skippers were studied from May through October 1982 in the 0.9- 
ha vegetable and flower garden where Lazri and Barrows (1984) in- 
vestigated flower visiting in Pieris rapae L. The garden is a community 
garden used in 1982 by about 146 gardeners, and it contains about 265 
species of entomophilous plants, including vegetables, ornamentals, 
herbs, wildflowers, and weeds. 

Flowers visited by the skippers and the relative numbers of skippers 
present at each species were noted during a total of 12 30-min mean- 
dering walks made through the garden twice each month in June, July, 
August, and September. The walks were made once every 2 weeks on 


VOLUME 39, NUMBER 4 301 


a clear day, every hour on the hour, from 0800 to 2000 h (EDT). At 
each skipper-frequented flowering plant or group of such plants, we 
made short (10 sec) counts to standardize the amount of time spent at 
a plant or group of plants. A total of 564 skippers of both species was 
counted during the entire census. 

To measure flower corolla lengths, we collected flowers in plastic 
bags and kept them moist until they could be examined. Dial calipers, 
accurate to 0.01 mm, were used to measure corollas (Lazri & Barrows, 
1984). We made a pollen reference collection from pollen collected in 
the study area. 

In studying possible flower constancy, frequency of flower use, and 
pollen deposition of skippers, we observed 22 foraging A. campestris 
and 60 foraging E. clarus. A stopwatch and tape recorder were used 
when needed. To discriminate focal individuals from other nearby 
skippers when they were common, we marked forewings of focal in- 
dividuals with small spots of enamel paint, which did not appear to 
affect their behavior. Forty additional skippers were each observed for 
10 min as they foraged at asteriad disk and ray florets. 

In examining possible pollen transport and deposition, we collected 
285 A. campestris and 77 E. clarus; 3 to 23 males and 5 to 25 females 
were taken from each VF VFS. Before it was captured, each skipper 
was followed as it visited two consecutive flower heads, extending its 
proboscis into a flower in each head for at least 1 sec. After it was 
netted, a skipper was paralyzed by carefully pinching the sides of its 
thorax between a thumb and forefinger and then placed into a glassine 
envelope on which relevant data were recorded. The enveloped skip- 
per was immediately put into an insulated bag filled with frozen cold 
packs. Within the hour, all skippers in the bag were put into a cooler 
filled with more frozen cold packs. At the end of a collecting day, the 
skippers were put into a freezer until they could be examined for pollen 
(Turnock et al., 1978). 

In searching for pollen on a skipper, we removed its legs and pro- 
boscis, placed them on a clean glass slide, and covered them with a 
drop of Permount® and a coverslip. The rest of the skipper was placed 
on a watchglass. Its proboscis, legs, body, glassine envelope, slide, and 
watchglass were examined for pollen under a compound microscope 
(up to 400 power), a dissecting microscope (up to 30 power), or both. 
Pollen adhering to the skipper’s labial palpi were included in its facial 
cavity count. Free floating pollen grains on the slide and watchglass 
and in the envelope were also counted. Adult skipper age was estimated 
to be young, middle-aged, or old, based on the amount of scale loss 
and wing tattering that was present on a skipper’s wings and body. A 
young skipper was one that was almost totally intact; a middle-aged 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


one had slight wing tattering and a few scales missing; and an old one 
had very tattered wings and many scales missing. 

Quantitative analyses were made with the Statistical Analysis System 
(SAS) computer package (Ray, 1982a, b). Pollen count and corolla 
depth values were log transformed to obtain homoscedastic data for 
the Duncan’s multiple range test (DMRT). Possible differences be- 
tween groups were analyzed with the t-test (TT) or paired t-test (PTT) 
corrected for heteroscedasticity when necessary, the Fisher exact prob- 
ability test (FEPT), and the Chi-square test (CST). Kendall’s rank cor- 
relation coefficient (KRCC) was used to test for significant correlations. 


RESULTS AND DISCUSSION 
Flowers Visited 


Atalopedes campestris visited 23 species of flowers (including two 
hybrids), in eight plant families (Table 1). Thirteen of these flowers 
(57%) were in Asteraceae, making it the most visited family. All but 
one of A. campestris’ eight very frequently visited flower species 
(VF VES) were asteriads. The other VF VFS was the dipsacaciad Sca- 
biosa atropurpurea which has florets that are morphologically similar 
to asteriad disk flowers. Atalopedes campestris also abundantly visited 
the asteriad Cosmos sulphureus, but since they did so for only 1 week, 
this flower was not classified as a VF VFS. 

Epargyreus clarus visited 28 species of flowers (including three hy- 
brids), in 16 plant families (Table 1). Eight of the flowers (29%) were 
asteriads. This skipper used only two VF VFS, both also VF VFS of A. 
campestris. Seven frequently visited flower species (F VFS) were used 
by E. clarus. Of these, Consolida orientalis and Dianthus barbatus 
were visited early in the season when few other flowers were in bloom. 
Epargyreus clarus visited Phaseolus vulgaris and Cucurbita sp. in mid- 
season when its VF VFS were commencing to bloom. All of the flowers 
visited by the skippers are introduced ones, except for Oenothera bien- 
nis, upon which only one E. clarus was seen and Eupatorium coeles- 
tinum which was visited by many A. campestris. 

Based on the censuses made during walks through the garden, we 
found three A. campestris adults in June, 40 in July, 65 in August, and 
176 in September on its VF VFS. These observed frequencies were 
significantly different from a hypothetical situation with equal monthly 
frequencies of 71 (the 4-month average) skippers (P < 0.001, CST). 
Twenty E. clarus visited their VF VFS in June; 34 in July; 72 in August, 
and 18 in September. These empirical frequencies were also different 
from a hypothetical situation with equal monthly frequencies (386) of 
skippers (P < 0.001, CST). These observed differences from equal 


VOLUME 39, NUMBER 4 303 


TABLE 1. Flowers visited by Atalopedes campestris (AC) and Epargyreus clarus 
(EC). Flower species are listed in systematic order by families (Bailey and Bailey, 1976) 
and alphabetical order by genera. 


Family Butterfly visitors 
Species, common name 

Liliaceae 

Allium vineale L., field garlic EC 

A. schoenoprasum L.., chives AC 
Amaryllidaceae 

Amaryllis belladonna L., belladonna lily EC 
Caryophyllaceae 

Dianthus barbatus L., sweet william AGES 
Ranunculaceae 

Consolida orientalis (J. Gray) Schrédinger, larkspur AC, EC 
Oxalidaceae 

Oxalis corniculata L., creeping oxalis AC 
Bassicaceae 

Rhaphanus sativus L., radish EG 
Fabaceae 

Phaseolus vulgaris L., snap bean EC 
Balsaminaceae 

Impatiens balsamina L., garden balsam EC 

Impatiens wallerana Hook., “Liegnitzia” Ee 
Violaceae 

Viola x wittrockiana Gams., garden pansy EC 
Lythraceae 

Lythrum salicaria L., purple or spiked loosestrife EC 
Onagraceae 

Oenothera biennis L., evening primrose EC 

O. fruticosa L., sundrops EC 
Boraginaceae 

Myosotis scorpioides L., true forget-me-not EC 
Laminaceae 

Lavandula sp., lavendar EC 

Mentha x piperita L., peppermint AC 

M. spicata L., spearmint AGLEE 

Ocimum basilicum L., common or sweet basil AC. EC 

Origanum vulgare L., marjoram or oregano AC, EC 
Solanaceae 

Capsicum sp., bell pepper AC 

Petunia x hybrida Hort. Vilm.-Andr., petunia EG 
Dipsacaceae 


Scabiosa atropurpurea L., sweet scabious AG EC 


304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Continued. 


Family Butterfly visitors 
Polemoniaceae 
Phlox paniculata L., phlox EC 
Asteraceae 
Ageratum houstonianum Mill., floss flower EC 
Aster cv., aster AC; EC 
Callistephus chinensis (L.) Nees., China aster AC? 
Centaurea cyanus L., bachelor’s button AC, EC 
Chrysanthemum leucanthemum L., ox-eye daisy AC, EC 
Chrysanthemum sp., chrysanthemum AC 
Cosmos bipinnatus Cav., cosmos AC, EC 
C. sulphureus Cav., cosmos AC 
Dahlia pinnata x coccinea Cav., dahlia AC,? EC 
Eupatorium céelestinum L., hardy ageratum AC,? EC 
Rudbeckia fulgida Ait., gloriosa daisy AC 
Tagetes erecta L., African marigold AC? 
T. patula L., French marigold AC? 
Zinnia elegans Jacq., zinnia AC,? EC 


4 Very frequently visited flower species. 


frequencies are expected due to seasonality of flowering and fluctuat- 
ing skipper population levels. 

The skippers used. VF VFS of different colors and similar shapes. 
Corolla tubes of many of these flowers were significantly different in 
length (Table 2). 


Flower Constancy 


Seventy-eight of the 82 observed skippers showed flower species 
constancy by visiting individual conspecific flowers or capitula twice 
in a row. This is significantly different from a hypothetical group in 
which by chance, 41 skippers visited conspecific flowers and 41 visited 
heterospecific flowers in sequence (P < 0.001, CST). All four of the 
skippers that visited heterospecific flowers were E. clarus foraging ear- 
ly in the season at their FVFS or IVFS (Dianthus barbatus and Ly- 
thrum salicaria, respectively). 

Further, 32 of the 82 skippers were observed as they made up to 14 
consecutive visits to flowers or capitula (Table 3). Fourteen of these 
skippers visited only 1 flower species; 14, 2 species; and four, 3 to 4 
species. Thus, the skippers tended to visit less than three flower species 
during a foraging bout. 

In asteriads, skippers preferred disk florets to ray florets. Forty ran- 
domly chosen A. campestris were each observed foraging for at least 
10 min on the flowers of Aster cv., Eupatorium coelestinum, Tagetes 


VOLUME 39, NUMBER 4 305 


TABLE 2. Corolla depths of very frequently visited flower species of Atalopedes 
campestris. Disc florets were measured for all flowers except Scabiosa atropurpurea for 
which regular florets were measured. Means followed by the same letter are not signifi- 
cantly different from one another (P < 0.05, DMRT). 


Corolla depth (mm) 


Flower species Mean = S.E., range, n 
Tagetes patula 26.74 + 0.214, 23.7—29.0, 34 
T. erecta 20.87 + 0.237, 17.7—25.0, 35 
Scabiosa atropurpurea 12.73 + 0.481, 8.0-18.0, 38b 
Zinnia elegans 12.04 + 0.350, 7.3-15.8, 26be 
Dahlia pinnata x coccinea MIES22=-70:320.0 7-4 10-4.a5be 
Callistephus chinensis 11.26 + 0.204, 9.5-15.0, 35c 
Aster spp. 7.67 + 0.145, 6.3-9.9, 85 
Eupatorium coelestinum 3540-0 IT 85.0) 35 


patula, and Zinnia elegans. All foraged significantly more (P < 0.05, 
FEPT) on the innermost mature disk flowers than on ray flowers when 
empirical data were compared to hypothetical cases in which skippers 
foraged at equal numbers on each of the two kinds of flowers. 


Stigma Contact 


All of the VF VFS of both skipper species have narrow corolla tubes 
with stigmas and anthers in positions that should promote proboscis 
contact as skippers imbibe nectar. Feeding skippers usually placed only 
their proboscis tips into corolla tubes, the remainders of their probos- 
cides bending above corolla tube openings. Some skippers feeding at 
flowers with longer corolla tubes, such as Tagetes erecta, T. patula, 
and the infrequently visited Consolida orientalis, Viola xX wittrock- 
iana, and Capsicum sp., occasionally pushed their proboscides deep 
into corolla tubes, possibly effecting stigma and anther contact with 
their “faces” and palpi. 


Pollen Transport 


From their VF VFS, 283 A. campestris carried 68.385 + 3.250 (0- 
357) pollen grains, and 59 E. clarus carried 45.05 + 8.760 (0-148) 
pollen grains. Atalopedes campestris carried the greatest mean amount 
of pollen from Eupatorium coelestinum, the only VF VFS that is native 
to the study-site region. The pollen loads that A. campestris carried 
from some species were significantly different from one another (Table 
4). Regarding their VF VFS, E. clarus carried significantly more Sca- 
biosa atropurpurea than Zinnia elegans pollen grains (P < 0.05, 
DMRT). 

Selected examples of significant differences in numbers of grains 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Sequential flower visits of individual skippers. AH, Ageratum houstonian- 
um; CA, Capsicum sp.; CC, Callistephus chinensis; CS, Cosmos sulphureus; DB, Dian- 
thus barbatus, DX, Dahlia pinnata x coccinea; EC, Eupatorium coelestinum; IW, Im- 
patiens wallerana; LS, Lythrum salicaria; SA, Scabiosa atropurpurea; TE, Tagetes erecta; 
TP, Tagetes patula; ZE, Zinnia elegans. 


Skipper number Sequence of flower visits 


Atalopedes campestris 


IL SA (14 times) 
2 ZE (10) 
3 ZE (6), SA (2), TE (3) 
4 TP (9) 
=) TE (4), EC (2), TE (2), EC (1), TE (1), EC (2) 
6 TP (5), DX (1), TP (6), CA (1) 
7 ZE (8), CC (1), ZE (2) 
8 ZE (5), SA (2), ZE (8) 
9 INES) SIAN Gl) aE) 
10 TE (6), EC (4), TE (1) 
It EC (10) 
2 ZE (11) 
Epargyreus clarus 
Ih ZE (10) 
2 ZE (1), LS (8), ZE (5) 
3 SA (1), DB (5), SA (3) 
4 ZE (1), DB (2), ZE (6) 
5) SA (10) 
6 DB (8) 
7 ZE (1), DB (4), ZE (5) 
8 ZE (11) 
9 ZE (11) 
10 ZE (5), SA (1), ZE (5) 
11 IW (4), ZE (2), SA (1), ZE (2) 
12 SA (18) 
13 ZE (10), IW (2) 
14 SA (11) 
15 SA (12) 
16 SA (5), CS (2), SA (2) 
17 SA (10) 
18 ZE (5), SA (1), AH (1), SA (4) 
19 ZE (4), SA (1), ZE (2), SA (2) 
20 ZE (13) 


carried by different skipper parts are listed in Table 5. Individual 
skippers carried significantly more pollen of these flowers in their facial 
cavities than on, or in, other structures, except for pollen of Tagetes 
patula. The most pollen any one A. campestris carried in its facial 
cavity (and in fact on, or in, any part) was 161 grains from Eupatorium 
coelestinum. For seven of their eight VF VFS, A. campestris carried 
the second largest amounts of pollen on their proboscides. Epargyreus 
clarus carried more pollen from Scabiosa atropurpurea and Zinnia 


VOLUME 39, NUMBER 4 307 


TABLE 4. Mean number of pollen grains carried by Atalopedes campestris and Epar- 
gyreus clarus from very frequently visited flower species and Cosmos sulfureus. Within 
a skipper species, means followed by the same letter are not significantly different from 
one another (P > 0.05, DMRT). 


Flower species Mean = SE, range, n 


Atalopedes campestris 


All very frequently visited flower species 68.35 + 3.250, 0-357, 283 
Cosmos sulphureus 135.83 + 15.662, 11-240, 16a 
Eupatorium coelestinum 116.22 + 9.298, 21-357, 45ab 
Aster cv. 91.55 + 9.350, 0-259, 42bc 
Dahlia pinnata x coccinea (3 D0 Ee la2. S=1665, 2c 
Zinnia elegans 59.04 + 4.042, 16-132, 48cd 
Callistephus chinensis 56:00) cen > 554-J9=139. Sede 
Tagetes patula 40.41 + 3.755, 8-142, 35de 
Scabiosa atropurpurea 37.29 + 4.222, 7-108, 35e 
Tagetes erecta 26.60 + 3.821, 0-103, 35 
Epargyreus clarus 
Both flower species 48.05 + 3.760, 0-148, 59 
Scabiosa atropurpurea 04.28 + 6.172, 0-135, 43 
Zinnia elegans So.06r= 4371) 9-143 2D 


elegans in their facial cavities than on, or in, any other parts. They 
carried significantly more Scabiosa atropurpurea pollen than Zinnia 
elegans pollen in their facial cavities (P < 0.001, TT). Epargyreus 
clarus from both flowers carried the second largest amounts of pollen 
on their proboscides. 

The significant differences in Table 5 indicate that skippers gener- 
ally carried more pollen anteriorly and less posteriorly. In order of 
decreasing amounts of pollen, E. clarus carried pollen in facial cavities, 
on proboscides, on hindlegs, on forelegs, in genital cavities, and on 
middle legs from Zinnia elegans and in facial cavities, on proboscides, 
forelegs, hindlegs, and middle legs and in genital cavities from Scabiosa 
atropurpurea. When mean numbers of pollen grains of two kinds of 
legs (or legs versus genital cavity) were compared, they were not usu- 
ally significantly different from one another. In contrast, pollen loads 
carried in skipper facial cavities and on proboscides were significantly 
greater than those carried by legs and genital cavities. Pollen of VF VFS 
was distributed similarly on A. campestris. 

The skippers usually carried more pollen of VF VFS on their pro- 
boscides or in their facial cavities compared to on, or in, other parts; 
this probably resulted from their more frequently putting their tongues 
rather than tips of their legs or other parts into flowers. Pollen from 
proboscides then builds up in facial cavities as skippers recoil their 
proboscides between flower visits. In cavities, pollen is likely to stick 


308 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 5. Selected examples of significant differences (P < 0.05, PTT) in the pollen 
loads carried between two body parts of an individual skipper from very frequently 
visited flower species. AC, Aster cv.; see Table 3 for other plant abbreviations. 


Compared parts Plants with significant differences 


Atalopedes campestris 


Facial cavity > proboscis AC, ZE 

Proboscis > hindleg AC, CC, DX, EC, TE, TP, ZE 

Hindleg > foreleg AS, TE 

Hindleg > middle leg AC, EC, SA, TE, ZE 

Foreleg > middle leg EC, SA, TE, TP, ZE 

Hindleg > genital cavity AC, EC, SA, ZE 

Foreleg > genital cavity AC, EC, SA, ZE 

Middle leg > genital cavity AC, SA, ZE 
Eparygreus clarus 

Facial cavity > proboscis SA 

Hindleg > foreleg ZE 

Hindleg > middle leg ZE 

Foreleg > middle leg SA, ZE 

Hindleg > genital cavity SA, ZE 

Foreleg > genital cavity SA, ZE 

Middle leg > genital cavity ZE 


to scales and other pollen already present and remain relatively un- 
disturbed. Skipper posture and movement on asteriad heads could also 
account for the pollen distribution on their bodies. On these capitula, 
they often have their heads over innermost, polliniferous disk florets 
and their thoraces and abdomens over outermost (female stage) disk 
florets and ray florets with little or no pollen. Further, skippers are 
likely to have smaller pollen loads on their legs and other more exposed 
parts due to pollen loss during locomotory and grooming behaviors. 
Some pollen was found in the genital cavity of at least one skipper 
collected from each of the VFVFS. This pollen was confined to the 
hairlike scales on females’ papillae analis and males’ valvae (Ehrlich, 
1960). The mean number of pollen grains borne in genital cavities for 
all VF VFS ranged from 0.17 + 0.171 (0-6, N = 85) grains of Scabiosa 
atropurpurea to 4.12 + 1.880 (0-15, N = 8) grains of Callistephus 
chinensis. Females of both skipper species carried significantly more 
pollen of these flowers, except for Dahlia pinnata x coccinea, in their 
genital cavities than males (P < 0.05, TT). Of the 41 skippers that 
carried pollen in their genital cavities, only eight were males. The most 
pollen any one female skipper carried in her genital cavity was 67 
pollen grains of Tagetes patula. The most pollen any one male skipper 
carried in his genital cavity was seven grains from Tagetes erecta. 
Pollen is likely to enter genital chambers when skippers touch pollen 
on flowers with their abdominal tips during foraging. Perhaps groom- 


VOLUME 39, NUMBER 4 309 


ing movements also cause pollen to enter genital chambers. Since the 
amount of pollen that a skipper carries might increase with its age, we 
examined our data for possible positive correlations between age and 
the pollen load of VF VFS on a skipper’s forelegs, middle legs, hindlegs, 
and proboscis, and in its facial and genital cavities. In A. campestris, 
age was positively correlated (P < 0.05) with the amount of pollen in 
facial cavities (8 flower species) and genital cavities (4), and on forelegs 
(1), hindlegs (1), and proboscides (5). In E. clarus, age was positively 
correlated with the amount of pollen in facial cavities (2 flower species) 
and on proboscides (2). Thus, pollen loads were generally not positively 
correlated with age, because out of a possible 48 correlations, only 13 
were found for A. campestris, and out of a possible 12 such correlations 
only four were found for E. clarus. 

In our study, the skippers visited Zinnia elegans more than other 
flowers. However, they carried more pollen from Eupatorium coeles- 
tinum and Scabiosa atropurpurea than from Zinnia elegans, which 
might not be expected (Heinrich & Raven, 1972). This finding might 
be due to Zinnia elegans having larger (55-u-diameter) and spinier 
pollen than the other two species which have 35-u-diameter pollen 
(Erdtman, 1966; Kapp, 1969). Pollen with a smoother exine surface 
adheres better to parts of Lepidoptera, such as tongues, than pollen 
with a spinier surface (Kislev et al., 1972). Further, pollen of smaller 
rather than larger diameter is generally picked up by a butterfly’s 
proboscis during feeding and is retained when its proboscis is recoiled 
and not in use (Levin & Berube, 1972). Besides skipper-visitation fre- 
quency and pollen size and surface characteristics of a particular flower 
species, the pollen load of a skipper is likely to be affected by many 
other variables which have not been studied. 

In conclusion, three lines of evidence suggest that the skippers were 
pollinators of their VF VFS. First, they were ordinarily constant to 
particular species during foraging bouts. Second, they transported pol- 
len. Finally, they were likely to contact stigmas frequently with their 
proboscides, since they mainly visited flowers with narrow, tubular 
corollas. They contacted some flowers with their facial cavities and 
may have contacted stigmas with their pollen-bearing legs and genital 
cavities, as well, because they sometimes walked over stigmas. 

However, two more important lines of evidence indicate that the 
skippers probably functioned mainly as nectar thieves. First, they most- 
ly foraged upon asteriads rather than upon other kinds of flowers and 
primarily probed innermost (male-stage) disk florets, tending not to 
contact female-stage florets with their more pollen-laden parts. Second 
and more importantly, the skippers carried pollen in loads that appear 
too small for efficient pollination. In the Colias-Phlox pollination sys- 


310 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


tem, Levin and Berube (1972) found that only 0.5% of the Colias- 
transported pollen, that was transferred to receptive stigmas, germi- 
nated and produced pollen tubes. In our study, 0.5% of the mean 
number of pollen grains from VFVFS that was carried by an entire 
individual skipper was always less than one grain. Making the liberal 
assumption that all pollen grains in the skippers are available for pol- 
lination and extrapolating from the Colias-Phlox system, we find that 
it would take an average of at least four flower visits for a skipper to 
deposit a pollen grain. If one considers an efficiently working capitu- 
lum to be one with many florets that can be pollinated by a single 
foraging insect (Burtt, 1961), capitula of the VF VFS are not efficiently 
working ones with regard to the skippers we studied. 

The butterflies-as-nectar-thieves hypothesis is not rejected by our 
observations on skippers and quantification of their pollen loads. Em- 
mel (1971) presents strong circumstantial evidence that the Ecuadorian 
skipper Perichares philetes dolores (Reakirt) is a pollinator of the or- 
chid Maxillaria “ontoglossom’” (which is not in Index Kewensis). How- 
ever, because he does not present direct evidence that this skipper is 
indeed a pollinator, his data do not reject the hypothesis. A further test 
of the hypothesis based on an extensive literature survey will be pre- 
sented in a future paper. 

The hypothesis that the skippers might indirectly increase pollina- 
tion and seed set of their thieved flowers remains to be tested. This 
increased pollination might occur because pollinators have to visit more 
flowers to obtain adequate resources from thief-depleted flowers com- 
pared to ones not depleted by thieves (Heinrich & Raven, 1972; Bar- 
rows, 1976). An alternative hypothesis to consider is that skippers some- 
how cause pollinators to forage less on patches of thieved flowers, 
thereby reducing pollination of these patches (McDade & Kinsman, 
1980; Roubik, 1982). 


ACKNOWLEDGMENTS 


We are grateful to the following persons who helped us with this study: R. S. Blanquet, 
F. M. Harrington, F. C. Thompson, L. Venables, and members of the Glover Park Garden 
Club. 


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Journal of the Lepidopterists’ Society 
39(4), 1985, 313-320 


PERMANENT TRAPS FOR MONITORING BUTTERFLY 
MIGRATION: TESTS IN FLORIDA, 1979-84 


THOMAS J. WALKER 


Department of Entomology and Nematology, University of Florida, 
Gainesville, Florida 32611 


ABSTRACT. Three models of a flight trap made principally of hardware cloth were 
tested at Gainesville, Florida. All models had a 6 m long central barrier of ¥% inch mesh 
hardware cloth. Butterflies encountering opposite sides of the barrier were trapped sep- 
arately, allowing calculation of net movement up or down the Florida peninsula. The 
most efficient model has a barrier 3.7 m high and a two-stage trapping superstructure of 
Y, inch hardware cloth. It catches 22-70% of migrant Phoebis sennae, Agraulis vanillae, 
and Urbanus proteus. 


Migrating butterflies characteristically fly in a straight line a few 
meters above the ground and rise and fly over obstacles rather than 
deviating laterally (Williams, 19380). Beginning in 1975, I have used 
stationary flight traps that intercept and trap migrant butterflies at 
Gainesville, Florida (Walker, 1978, 1980; Walker & Riordan, 1981). 
My first traps were made of polyester, which ripped in strong winds 
and deteriorated in sunlight. They consequently required frequent re- 
pair and annual replacement. Furthermore, they lost about 90% of the 
migrants they intercepted. 

In this paper I describe the development of a hardware-cloth trap 
that will work for years without repair and that promises, with speci- 
fied improvements, to catch more than 70% of the migrants that en- 
counter it. 


THE TRAPS 


Three models of permanent flight traps were tested. All resembled 
the polyester traps in having a 6 m long central barrier oriented ENE- 
WSW (perpendicular to the Florida peninsula) and a holding device 
at either end. All kept the butterflies that had encountered the barrier 
from the migratory direction +90° separate from those that had en- 
countered it from the opposite direction +90°. 

Model #1. The first trap (Fig. 1, right) was constructed during 
February 1979 in a pasture with scattered trees, northwest of Gaines- 
ville (NW4, sec. 31, tp. T9S, RI9E). The central barrier was of % inch 
hardware cloth attached to three pressure-treated “4x 4’ posts (i.e., 
9 x 9 cm). The roof, also of % inch hardware cloth, was 1.2 m from 
ridge to eave and was fastened laterally and medially to treated “2 x 4’s” 
(4 x 9 cm). Its ridge slanted upward from the center post (3.4 m high) 
to either end post (4.0 m), in imitation of a polyester trap (see fig. 1 
of Walker, 1978). The roof sloped 30° toward its eaves. Migrant but- 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Models #8 (left) and #1 (right) of a permanent flight trap for migrating 
butterflies. 


terflies were to encounter the central barrier, be detained between the 
roof and the barrier, and work their way upward to the nearest end. 
There they were to continue upward through an 8 x 24 cm opening, 
through an immovable hardware cloth “valve,” and into a holding cage 
of plywood and % inch hardware cloth. Watching migrants encounter 
model #1, I discovered that most individuals shunned the offered 
openings and instead flew out and over the roof or around the end 
“wall” (i.e., panels of % inch hardware cloth that extended 1.2 m from 
either end of the central barrier and perpendicular to it). 

Model #2. During August of 1983 I constructed a second trap im- 
mediately ENE of the first. It differed from model #1 in having a 18 
cm slot along the entire upper edge of each roof panel. These slots 
gave access to a longitudinally partitioned 6.0 x 0.4 x 0.4 m duct of 
Y, inch hardware cloth that prevented the butterflies’ escaping as they 
worked their way to either end of the trap, through hardware cloth 
valves and into holding cages. The central barrier was rectilinear and 
3.7 m high. The roof ridge was made straight and the roof slope was 
reduced to 15°—making the eaves 3.4 m high. Although model #2 
caught substantially higher proportions of migrants than model #1, 
most migrants were hesitant to fly through the 18 cm slots and would, 
instead, hover under the roof and eventually escape. 

Model #3. During February 1984 I greatly improved access to the 
longitudinal duct, thereby converting model #2 to model #3. The 
width of the roof slots was increased more than threefold to 45 cm and 
a sharply sloping upper roof of % inch hardware cloth was interposed 
between the duct and each main roof (Fig. 1, left; Fig. 2). The hard- 


VOLUME 39, NUMBER 4 B15 












A 


partitioned holding 


cage 


partitioned holding cage —duct and duct baffle 
with doors for 
removing buiterflies 7— upper roof 


-— main root 


Ni 2x 4 cross- 


piece with 

2 c adjoining 
valves —~ upper edges 
of roof panels 


/ 


0.5 m slot 















3.3 m 


—central barrier 


cross-section 0.2 m 
from either end 


— central support pole 
(supporting joined 
roof panels) 





I 
flat steel for 


securing poles 


— roof panel support pole, 
1° conduit 


Fic. 2. Model #8 permanent flight trap: A lod i d 
p: A, exploded diagram; B, 
(Drawings by S. A. Wineriter) gr Grass “section: 


316 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Migration of four species of butterflies as revealed by 6 m, permanent flight 
traps at Gainesville, Florida, 1979-1984. 


2 Net northward spring migration* Net southward fall migration> 
ra ee Eee 
ean P.sen. A.van. J.coenia  U. prot. P. sen. A.van. J.coenia_ U. prot. 
Model #1 
1979 0 1 185 0 157 22 2 13 
1980 3 0 13 0 69 12 i 9) 
1981 2 0 44 0 263 26 6 i 
1982 a 2 85 0 126 15 22. 24 
1983 2 0 4 0 54 q = 0 
1984 ] ] 27 0) 92 10 =) 0 
Model #2 
1983 — — — — 86 157 21 ray9) 
Model #3 
1984 6 6 252 =a 548 326 62 531 
Sum 21 10 610 = 1395 575 110 629 
Consistency° 96 92 96 33 91 94 72 99 


* Number trapped on south side of barrier minus number trapped on north side of barrier (1 March to 22 May). 
> Number trapped on north side of barrier minus number trapped on south side of barrier (1 Sep. to 30 Nov.). 
¢ Percent of total trapped that were flying in the migratory direction (viz. southward in the fall, northward in spring). 


ware cloth of the 50 cm upper roof extended as a baffle 25 cm into 
the duct, thereby impeding the escape of migrants from the duct (Fig. 
2B). (Building a #3 trap is described in the appendix.) 


THE CATCHES 


At least seven species of butterflies migrate southward through 
Gainesville each fall: Phoebis sennae (L.), Agraulis vanillae (L.), Ju- 
nonia coenia Hibner, Urbanus proteus (L.), Panoquina ocola (Ed- 
wards), Lerema accius (J. E. Smith), and Eurema lisa (Boisduval & 
LeConte) (Walker, 1978, 1980, 1985). Only the first four will be dealt 
with here, because they were captured in the largest numbers. 

As reported previously (Walker, 1980), the direction of net move- 
ment of these species at Gainesville is down the peninsula in the fall 
and, for the first three species, toward Georgia in the spring (Table 1). 
Net numbers trapped flying northward in spring (1 March to 22 May) 
for the six years varied from —1 for U. proteus (viz., 1 northward, 2 
southward) to 610 for J. coenia. Net numbers trapped flying southward 
in fall (1 Sep. to 30 Nov.) varied from 110 for J. coenia to 1395 for P. 
sennae. With the exception of U. proteus in spring and P. coenia in 
fall, more than 90% of migrants trapped were captured flying in the 
seasonally appropriate direction (Table 1). 

Trapping efficiency of models #1 and 3 was studied during October 
1984. During the five observation periods of 3 hours or more, model 


VOLUME 39, NUMBER 4 oly 


TABLE 2. Absolute trapping efficiency of model #3 of a permanent trap for sampling 
migrating butterflies. 


P. sennae A. vanillae U. proteus 
Capt./ Capt. / Capt./ 

Date (1984) Time (EDT) cand.?* % cand. % cand. % 

4 Oct. 1251-1551 16/25 64 6/11 50 Sy PAS) 60 

3 Oct. 0917-1217 14/17 82 3/7 43 11/23 48 

5 Oct. 1306-1606 8/12 67 0/6 0 9/19 47 

11 Oct. 1288-1600 12/18 67 4/13 31 3/10 30 

12 Oct. 1100-1400 8/24 33 BUS 33 3/7 43 

All observations 58/96 60 18/52 35 41/84 49 
95% C.I.> 49-70 22-50 38-60 


*Number of migrants captured during observation period/number of candidate migrants (i.e., southward flying 
individuals that would have flown over the 6-m, ENE-WSW line at the base of the traps central barrier had the trap 
not been in place). 

> Based on binomial distribution. 


#3 caught an average of 60% of candidate P. sennae, 35% of A. vanilla, 
and 49% of U. proteus (Table 2). Model #1 caught 13% of candidate 
P. sennae (18 of 98) but O of 44 A. vanillae and 0 of 55 U. proteus. 

Because the traps sample adjacent 6 m cross sections of migrants, it 
is likely that season-long differences in their catches are due principally 
to differences in trapping efficiency and that differences in numbers 
of potential captives are minor or lacking. Confirming this conjecture 
is the fact that numbers of P. sennae and A. vanillae observed during 
15+ hours of watching were 98 and 44 for model #1 and 96 and 52 
for model #8. (Numbers of U. proteus were more discrepant for the 
two traps, 55 and 84, but these butterflies are relatively small, dark, 
and fast, making it likely that some escaped notice—which, in turn, 
makes it likely that 49% overestimates the proportion of this species 
trapped.) Table 3 compares catches of models #2 vs. 1 during fall of 
1988 and catches of models #8 vs. 1 during all of 1984. 

By using the absolute trapping efficiencies in Table 2 and the relative 
trapping efficiencies in Table 3, the numbers of fall migrants in Table 
1 were converted to estimates of total fall migration across each ENE- 
WSW meter (Table 4). (All traps were oriented ENE-WSW—perpen- 
dicular to the axis of the Florida peninsula.) 


DISCUSSION 


Further improvements. The model #8 flight trap caught far higher 
proportions of the migrant butterflies that encountered it than did 
earlier polyester or hardware cloth traps (Table 3). However, its ab- 
solute efficiency was still less than 70% (Table 2). Two easy-to-make 
changes promise to improve its performance substantially. The first 


318 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Relative trapping efficiency of models #1, 2, and 3 of a permanent trap 
for migrating butterflies. 


Model 2 vs. 1 Model 3 vs. 1 Model 8 vs. 2 
Species of es eee EE aS ae 
migrant Numbers* Ratio Numbers? Ratio Ratio* 
P. sennae 86 vs. 54 1.6 504 vs. 93 6.0 3.7 
A. vanillae 157 vs. 7 22.4 332 vs. 11 30.2 1.3 
J. coenia PI ws, = ll — 314 vs. 24 13.1 i 
U. proteus 55 vs. O — 530 vs. O — ? 


* Net numbers of migrants caught by models 2 and 1 during fall 1983. 
> Net numbers of migrants caught by models 3 and 1 during spring and fall 1984. 
© Calculated by using model 1 as the standard. 


change concerns the fact that some migrants refused to fly upward 
into the longitudinal duct. The refusal of some of these migrants prob- 
ably resulted from their view of the sky being partially blocked by 6 
m of 2x4 that supported the duct. A less sky-blocking support (e.g., a 
3 x 3 cm steel angle) should be substituted. The second change con- 
cerns the fact that most of the migrants that escaped did so by flying 
around the end walls. (Specifically, 67 of the 115 escapees in Table 2 
left the trap within 10 seconds by flying around the end wall.) The 
end walls could be extended to 2.4 m making lateral escape much less 
likely. 

Uses. Permanent flight traps can monitor butterfly migrations con- 
tinually, and they can provide information about migrations so sparse 
that they cannot be directly observed. The data in Tables 1 and 4 (and 
unpublished data on other species) illustrate these uses. Permanent 
flight traps also provide a convenient means of collecting large numbers 
of live migrants for studies of morphology, physiology, sex ratios, mat- 
ing status, behavior, etc. 

Traps with other uses. The great improvement in efficiency of the 
model #8 over the model #1, which copied the design features of the 
original polyester trap (Walker, 1978), suggests that a much improved, 
portable, polyester trap might be made by copying the design features 


TABLE 4. Fall migration (net no. flying southward across each ENE-WSW meter) as 
estimated by permanent flight traps, Gainesville, Florida, 1979-1984. (Numbers captured 
are in Table 1; trapping efficiencies based on Tables 2 and 3. Estimates for 1983 and 
1984 are from catches of models #2 and 3, respectively.) 


Year 


Species 1979 1980 1981 1982 1983 1984 
P. sennae 262 115 438 210 88 152 
A. vanillae 316 173 374 216 97 155 


U. proteus —_ — — — — 181 


VOLUME 39, NUMBER 4 319 


of the model #3 permanent trap. Furthermore, traps half as long should 
catch much larger numbers of migrants than did the original 6 m 
polyester traps. (A similar shortening is also an option for permanent 
traps and would reduce costs for materials ca. 30%.) 

An important limitation for all flight traps yet used to study butterfly 
migration is that they distinguish migratory directions only crudely. 
This limitation could be overcome by constructing an octagonal trap 
having eight identical openings leading to eight holding cages, thereby 
separating migratory directions at 45° intervals rather than the 180° 
intervals of the present traps. 

The permanent traps built thus far capture migrants alive and, 
therefore, require daily servicing. Traps could be run at remote loca- 
tions, or at near locations with reduced service time, if the holding 
cages were modified to kill and preserve the migrants captured. For 
example, dichlorvos-impregnated plastic could be used to cause the 
captives to drop into containers of dilute formalin. 

Finally, devices could be substituted for the holding cages that would 
automatically mark the butterflies with fluorescent pink paint and al- 
low them to continue their migratory flights—to be caught, perhaps, 
by downstream traps. (If such devices seem far-fetched, see Wolf and 
Stimmann, 1972.) 


ACKNOWLEDGMENTS 


I thank T. G. Forrest, J. E. Lloyd, and S. A. Wineriter for constructively criticizing 
the manuscript. Susan A. Wineriter also contributed by measuring the efficiency of the 
traps (Table 2) and by artwork (Fig. 2). Florida Agricultural Experiment Station Journal 
Series No. 6091. 


LITERATURE CITED 


WALKER, T. J. 1978. Migration and re-migration of butterflies through north peninsular 
Florida: Quantification with Malaise traps. J. Lepid. Soc. 32:178-190. 

1980. Migrating Lepidoptera: Are butterflies better than moths? Pages 79-98 

in Insect behavioral ecology symposium, ‘79. Fla. Entomol. 63:1-111. 

1985. Butterfly migration in the boundary layer. Pages 704-723 in Migrations: 
Mechanisms and adaptive significance. Contrib. Mar. Sci. 27 (Suppl.). 

WALKER, T. J. & A. J. RIORDAN. 1981. Butterfly migration: Are synoptic scale wind 
systems important? Ecol. Entomol. 6:433—440. 

WILLIAMS, C. B. 1930. The migration of butterflies. Oliver and Boyd, Edinburgh. 473 








Pp. 
Wo LF, W. W. & M. W. STIMMANN. 1972. An automatic method of marking cabbage 
looper moths for release-recovery identification. J. Econ. Entomol. 65:719-722. 


APPENDIX 


This appendix describes the main steps in building a model #3 permanent flight trap. 
It omits details that can be improvised by anyone with experience in light construction. 
The present model #8 was built by modifying a model #2, but the following steps 
describe how to build one from scratch. Materials for one trap now cost ca. $500. 


320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1. Central supports. Lay out a 6 m line perpendicular to the migratory direction. At 
each end and at the center of the line set a post (e.g., an 18’ treated 4x4) so that 4.4 m 
extends vertically from the ground. Connect the posts at 3.7 m with treated 2 4’s (to 
which the main roof panels will be attached). 

2. Superstructure. Prepare a support for the duct by attaching %,” x 1” = 0.4 m cross 
pes. of flat steel at the ends and at 1.5 m intervals along one flat surface of a 6.0 m pc. 
of % x 1% x 1%" steel angle. Attach the steel angle, cross pcs. up, to the tops of the 
main posts. Affix a 0.7 m vertical support for the steel angle midway between each pair 
of main posts. Install two 3.0 x 0.7 m vertical partitions of % inch hardware cloth, 
attaching the top edges to the steel angle, the ends to the posts, and the bottom edges to 
the 24 cross pes. Make a three-sided square duct by bending lengthwise a 6.0 x 1.2 m 
pe. of % inch hardware cloth at 0.4 and 0.8 m. Invert the duct over the duct support 
and attach a 6.0 x 0.75 m pe. of % inch hardware cloth to each lower edge of the duct 
in such a fashion that the lower 50 cm of width can become upper roof and the upper 
25 cm of width can become duct baffle (Fig. 2). 

3. Main roof panels. Build four roof panel frames of treated wood and steel tubing, 
each consisting of a 2 x 4 x 1.7 m (outer rafter; make 2.5 m if end wall is to be 2.4 m), 
al x 4(=2 x 9cm) x 3.0 m (upper edge), a 1 x 4 x 1.7 m (inner rafter), and a 3.0 
m pe. of %” electrical conduit (lower edge). Cut upper ends of rafters at 75°. Attach a 
1.2 x 3.0 m pe. of % inch hardware cloth to each roof panel with one edge riveted to 
the conduit, leaving a 0.5 m slot between the hardware cloth and the upper edge of the 
panel frame. Attach each roof panel by its upper edge to one of the 2X4 cross pcs. 
Support the rafters at ca. 1.2 m with poles that position the eaves at 3.38 m. (Make poles 
of 2 pes. of 1” electrical conduit joined by driving them over opposite ends of a short pc. 
of %” galv. pipe.) Bolt together the inner rafters of adjacent roof panels. Attach the lower 
edge of the secondary roof to the upper edge of the main roof. 

4. Central barrier and ends. Attach the central barrier of three 6.0 x 1.2 m pes. of 
% inch hardware cloth to the main support posts. Close the ends of the duct and the 
secondary roof with %4” hardware cloth. Make the end walls by attaching 1.2 m wide 
pes. of %” hardware cloth to the end posts, the outer rafters of the roof panels, and the 
roof support poles. (If the end walls are to be 2.4 m wide, install another pole 1.2 m 
beyond each existing end-rafter support pole.) 

5. Attachments. Construct two 4.4 m ladders using treated 2x 4’s as side pieces and 
1” electrical conduit as rungs. Install one ladder 0.3 m away from each end post. At the 
top of each ladder secure a safety loop of % x 1” aluminum (to enable one to use both 
hands in servicing the trap). Make hardware cloth valves by appropriately cutting 15 x 
30 cm areas on each side of each end of the top of the duct. Build two partitioned holding 
cages that will fit over the valves at either end of the duct. Make the doors to the chambers 
of the holding cages so that they will stay open as butterflies are removed. Install the 
holding cages—and wait for migrants. 


ADDENDUM 


During March 1985 the model #1 trap was razed and in its place an improved model 
#3 trap (i.e., a model #4 trap) was built using the directions given above—except that 
the main roof was made horizontal, thereby, simplifying construction and elevating the 
duct by 11 cm. The end walls extended 2.4 m from the central barrier. During the period 
10 Apr to 29 May 1985, the net numbers of J. coenia trapped flying northward were 
216 for the model #3 and 302 for the model #4 trap, translating into a 40% improvement 
in catch. 

For the first time Vanessa virginiensis Drury was identified as a spring migrant, with 
11 trapped flying northward and 2 flying southward (chi-square = 6.23; P < 0.05). 


Journal of the Lepidopterists’ Society 
39(4), 1985, 321-327 


ADULT NOCTUIDAE FEEDING ON APHID HONEYDEW AND 
A DISCUSSION OF HONEYDEW FEEDING 
BY ADULT LEPIDOPTERA’ 


JAMES B. JOHNSON AND MICHAEL P. STAFFORD 


Department of Plant, Soil and Entomological Sciences, 
University of Idaho, Moscow, Idaho 83843 


ABSTRACT. Adult Aseptis characta (Grote) and Rhynchagrotis exertistigma (Mor- 
rison) (Lepidoptera: Noctuidae) were observed feeding on honeydew produced by Zyxa- 
phis canae (Williams) (Homoptera: Aphididae) on basin big sagebrush, Artemisia t. 
tridentata Nuttall, in south-central Idaho. The feeding behavior is described. The lack 
of similar reports among other Lepidoptera, excluding the Lycaenidae, is discussed. 
Means by which the moths could locate honeydew sources and the adaptive value of this 
ability in specific situations are considered. 


On 22 July 1983, a male Aseptis characta (Grote) and a female 
Rhynchagrotis exertistigma (Morrison) (Lepidoptera: Noctuidae) were 
observed feeding on the honeydew of Zyxaphis canae (Williams) (Ho- 
moptera: Aphididae) on basin big sagebrush, Artemisia t. tridentata 
Nuttall. This observation was made in a canyon 6 miles SSW of Howe, 
Butte Co., Idaho. The day was moderately cool (about 22°C) and cloudy, 
with a light rain falling. Despite the weather, there was a moderate 
amount of insect activity. Numerous aphids, flies and wasps, especially 
Ichneumonidae, were seen on the sagebrush, Artemisia spp. 

Each moth located an aggregation of aphids and moved its proboscis 
from aphid to aphid within that aggregation. Then, each probed with 
its proboscis and if necessary, walked a short distance to locate another 
group of aphids. The behavior was similar to locating composite inflo- 
rescences and probing individual florets. 

Feeding on homopterous honeydew is a logical extension of the typ- 
ical nectar-feeding habit of adult Lepidoptera. Both are aqueous so- 
lutions containing carbohydrates and amino acids, plus a variety of 
minerals, lipids, organic acids and vitamins (Hagen, 1958; Auclair, 
1968; Strong, 1963; Baker, 1977; Baker & Baker, 1979). Nectars may 
also contain potentially toxic compounds, e.g. alkaloids and glycosides 
(Baker, 1977; Baker & Baker, 1979). There are no reports of such 
potentially toxic compounds in honeydews. Therefore, it should be 
possible for many adult Lepidoptera to exploit this alternative food 
source. However, few accounts of honeydew feeding by adult Lepi- 
doptera could be located and all involved Lycaenidae (Bingham, 1907; 
Lamborn, 1914; Roepke, 1918; Farquharson, 1922; Balduf, 1939; Hin- 


' Published with the approval of the director of the Idaho Agricultural Experiment Station as Research Paper No. 
8475. 


322 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ton, 1951; Gilbert, 1976; Orsak, 1977; Henning, 1983). Species in some 
lycanenid genera are not known to visit floral nectaries, so homopteran 
honeydew may be their primary source of nutrients (Cottrell, 1984). 

All of the earlier reports of adult Lepidoptera feeding on homop- 
terous honeydew involved Lycaenidae. The family includes many, tax- 
onomically diverse species in which the larvae are predaceous on Ho- 
moptera, associated with ants in some way or both (Balduf, 1939; 
Clausen, 1940; Hinton, 1951; Henning, 1983). Species with homopter- 
ophagous or myrmecophilous larvae commonly oviposit near aggre- 
gations of Homoptera which serve as prey or indicators of an area 
where hosts are likely to be found, respectively. Therefore, it is not 
surprising that they have evolved the habit of exploiting honeydew as 
a readily available source of adult food. 

There is no evidence that A. characta or R. exertistigma would be 
expected to locate aphid aggregations for any purpose, other than as a 
source of honeydew. This implies that exploiting this alternative food 
source should be possible for many Lepidoptera, in addition to the 
entomophagous Lycaenidae. The paucity of observations of this be- 
havior in other groups could be due to Lepidoptera being less conspic- 
uous when feeding on honeydew than when feeding on nectar. Ento- 
mophilous flowers are usually prominently displayed so that they may 
be more readily located by their insect pollinators (Jensen & Salisbury, 
1972). Most aphid aggregations, on the other hand, are relatively in- 
conspicuous and may be effectively concealed, thus, also concealing 
any visitors. However, since many Lepidoptera are relatively large and 
easily observed, it seems unlikely that this behavior would not have 
been reported more frequently, if it were common. So, it seems prob- 
able that adult Lepidoptera, other than Lycaenidae, rarely consume 
honeydew. 

At this point two questions arise: 1) Why is honeydew feeding un- 
usual among adult Lepidoptera, other than Lycaenidae?; and 2) Why 
did it occur in the situation described earlier? Possible answers for both 
questions will be discussed. 

While adult Lepidoptera are generally regarded as nectar-feeding 
insects, they actually display considerable flexibility in their feeding 
behavior. “Puddling,” in the broad sense, includes feeding at the mar- 
gins of puddles, etc. and on urine, dung and carrion (Arms, Feeny & 
Lederhouse, 1974; Downes, 1973); it is common among adult Lepi- 
doptera. Some Lepidoptera are also known to feed on fluid from the 
eyes of mammals and mammalian skin secretions, including sweat, and 
blood flowing from wounds (Banziger, 1971; Buttiker, 1959, 1962, 1964). 
Still more specialized is behavior of the SE Asian noctuid Calyptra 
eustrigata (Hmps.), which moth uses its proboscis to pierce the skin of 


VOLUME 39, NUMBER 4 320 


large mammals to obtain blood meals (Banziger, 1971, 1975). Water 
and amino acids were considered to be the key nutrients acquired from 
these atypical food sources, but in at least some cases, the acquisition 
of sodium may also be of great importance (Arms, Feeny & Leder- 
house, 1974). The two latter nutrients could be deficient in specific 
nectars, therefore, these foods could be important supp!ements to diets 
that consist largely of nectar. In this situation, it is not surprising that 
Lepidoptera have evolved the habit of exploiting these food sources. 

Still, some adult Lepidoptera regularly consume atypical foods that 
would seem more suitable as substitutes for nectar, than as dietary 
supplements. African and Asian Sphingidae of the genus Acherontia, 
commonly enter nests of wild and domestic bees to consume stored 
honey (Balduf, 1939). Their probosces are recurved apically and rel- 
atively short, indicating that they may be specialized for feeding in 
this manner. Banziger (1970) discusses a Malayan noctuid, Calyptra 
thalictri (Bkh.), that uses its similarly structured proboscis to pierce 
fruit. (This behavior is assumed to be ancestral to the skin-piercing, 
blood-sucking habit of C. eustrigata.) In these two cases, the high 
concentrations of sugars in the foods suggest that carbohydrates may 
be among the nutrients sought by the moths. The high sugar concen- 
trations and the large volumes of food available in these situations seem 
to be plausible reasons for the moths evolving these unusual feeding 
habits. However, the possibility of some other nutrient(s) being of pri- 
mary importance to the moths cannot be totally discounted. 

Though few in number, these examples of highly modified feeding 
behaviors, which are apparently directed primarily to the acquisition 
of carbohydrates, are important. They make it more surprising that 
adult Lepidoptera do not consume homopterous honeydew more reg- 
ularly. Intuitively, it would seem that there must be some factors which 
have tended to restrict their exploitation of this food source. 

The Ditrysia, the dominant and most advanced suborder of the 
Lepidoptera, seem to have radiated ecologically and taxonomically, in 
synchrony with the Angiospermae (Common, 1970, 1975). With a co- 
evolutionary history that extends back about 100 million years to the 
mid-Cretaceous (Powell, 1980), it is not surprising that these Lepidop- 
tera have, in general, evolved the specialized ability to efficiently locate 
floral nectar sources using visual and olfactory cues (Brantjes, 1976). 

An aggregation of aphids would certainly not present visual stimuli 
like those of a flower or inflorescence. Nor would there be any stimulus 
analogous to floral odor to guide Lepidoptera to a source of honeydew. 
Yet, the occurrence of Lepidoptera at sap flows on injured trees and 
the successful use of molasses bait traps, demonstrate that some Lepi- 
doptera are capable of locating food sources using only olfactory cues 


324 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


other than floral odors. Therefore, it is possible that some Lepidoptera 
could locate honeydew sources, perhaps in a manner similar to that 
used by Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae). 
Adult C. carnea orient anemotactically to indole acetaldehyde, a 
breakdown product of the amino acid tryptophan which occurs in some 
honeydews (Hagen, Greany, Sawall & Tassan, 1976; van Emden & 
Hagen, 1976). So, the question remains, why is a convenient food 
source apparently underexploited by adult Lepidoptera? 

Competition with ants and other common honeydew-feeding insects 
seems likely to be an important factor. Unlike floral nectar, which is 
often protected by morphological and chemical systems which restrict 
access to specific groups of visitors (Grant & Grant, 1965; Heinrich, 
1970; Feinsinger, 1983), honeydew is usually exposed and freely ac- 
cessible to many visitors. Therefore, the honeydew-feeding niche would 
be more likely to be dominated by groups like the ants, which are 
abundant, aggressive, effectively search plant surfaces and are less spe- 
cialized in their feeding habits. 

The generally ready and dependable availability of flowers during 
the seasons of adult Lepidoptera activity may be equally important. 
Aphid populations are prone to sudden, dramatic increases, e.g. pest 
outbreaks, and decreases, e.g. following an Entomophthora sp. epizo- 
otic (Hagen, 1976) or a period of extreme heat (Neuenschwander, 
Hagen & Smith, 1975). This less stable situation would seem poorly 
suited to relatively short-lived, pro-oogenic species, like most Lepidop- 
tera (Chapman, 1982), which must rather quickly locate carbohydrate 
sources and deposit large numbers of already mature eggs. The influ- 
ence of dependably available nectar is indirectly supported by the 
circumstances that existed at the time this observation was made. 

In the high desert of south-central Idaho, mid-July is normally a 
time of transition, as the late spring flowers, e.g. desert paintbrush, 
Castilleja chromosa A. Nelson, globe mallow, Sphaeralcea munroana 
(Douglas) Spach ex Gray, and Chaenactis douglasii (Hooker) Hooker 
and Arnott, are passing and the summer flowers, e.g. rabbitbrush, 
Chrysothamnus spp. and horsebrush, Tetradymia spp. are coming into 
bloom. Sagebrush species in this area begin flowering in early fall and 
produce little, if any, nectar since they are anemophilous (Stebbins, 
1974). However, in July 1983, an atypical, prolonged period of cool, 
rainy weather seemed to substantially delay the bloom of the summer 
flowering shrubs in this area, leading to a temporary, but acute, short- 
age of nectar sources. 

Prolonged periods of moderately cool weather are also known to 
produce “aphid years.” This is believed to be due to the temperature 
remaining above the developmental threshold of the aphid species, but 


VOLUME 39, NUMBER 4 320 


below that of their predators and parasites (Neuenschwander, Hagen 
& Smith, 1975). This situation can lead to a rapid increase in the aphid 
population. An increase of this type would have increased the avail- 
ability of honeydew, thus, increasing the likelihood that the moths 
would discover this alternate food. Since it remained cool and rainy at 
the time this observation was made, the weather may have severely 
limited the moths’ abilities to search for and exploit the few nectar 
sources that were available. It seems likely that some combination of 
these factors induced the moths’ atypical behavior. 

So, honeydew feeding by adult Lepidoptera, other than entomopha- 
gous Lycaenidae, appears to be uncommon. If it were generally to 
occur only under circumstances similar to those outlined above, it would 
explain the scarcity of reports of this intuitively logical behavior. Still, 
under specific conditions, the ability to efficiently locate and consume 
honeydew could be important to the survival of many Lepidoptera. 

These observations were made in conjunction with work conducted 
for the Idaho National Engineering Laboratory Radioecology and 
Ecology Programs sponsored by the Office of Health and Evironmental 
Research, United States Department of Energy, under contract number 
DE-AM07-811D12210. 


ACKNOWLEDGMENTS 


We wish to thank Drs. John Rawlins, Department of Zoology, University of Texas, 
Austin, TX 78712 and John G. Franclemont, Entomology Department, Cornell Univer- 
sity, Ithaca, NY 14853 for identifying the Noctuidae. We are grateful to Dr. David J. 
Voegtlin, Illinois Natural History Survey, Natural Resources Building, Urbana, IL 61853 
for identifying Zyxaphis canae (Williams). We also wish to ackriowledge Dr. Jerry A. 
Powell, Department of Entomological Sciences, University of California, Berkeley, CA 
94720 for supporting our belief that this phenomenon has not been reported previously 
among moths and Dr. Richard A. Arnold, 50 Cleaveland Rd. #8, Pleasant Hill, CA 94523 
for reviewing the manuscript. 


LITERATURE CITED 


ARMS, K., P. FEENY & R. C. LEDERHOUSE. 1974. Sodium: Stimulus for puddling be- 
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AUCLAIR, J. L. 1963. Aphid feeding and nutrition. Ann. Rev. Entomol. 17:439—490. 

BAKER, H. G. 1977. Non-sugar constituents of nectar. Aphidologie 8:349-356. 

BAKER, H. G. & I. BAKER. 1979. Starch in angiosperm pollen grains and its evolutionary 
significance. Amer. J. Bot. 66:591—600. 

BALDUF, W. V. 1939. The bionomics of entomophagous insects. Pt. Il. John S. Swift 
Co., St. Louis. 384 pp. 

BANZIGER, H. 1970. The piercing mechanism of the fruit-piercing moth Calpe (Calyp- 
tra) thalictri Bkh. (Noctuidae) with reference to the skin-piercing moth C. eustrigata 
Hmps. Acta Trop. 27:54-88. 

1971. Bloodsucking moths of Malaya. Fauna 1:5-16. 

1975. Skin-piercing bloodsucking moths I: Ecological and ethological studies 
on Calpe eustrigata (Lepid. Noctuidae). Acta Trop. 32:125-144. 

BINGHAM, C. T. 1907. The fauna of British India including Ceylon and Burma, but- 
terflies. Vol. II, Lycaenidae. London. 480 pp. 








326 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BRANTJES, N. B. M. 1976. Senses involved in the visiting of flowers by Cucullia um- 
bratica (Noctuidae, Lepidoptera). Entomol. Exp. & Appl. 20:1-7. 

BUTTIKER, W. 1959. Observations on feeding habits of adult Westermanniinae (Lepid., 
Noctuidae) in Cambodia. Acta Trop. 16:356-361. 

1962. Notes on two species of Westermanniinae (Lepidoptera: Noctuidae) from 

Cambodia. Proc. Roy. Entomol. Soc. London (B) 31:73-76. 

1964. New observations on eye-frequenting Lepidoptera from S.E. Asia. Verh. 
Natur. Ges. Basel. 75:231-236. 

CHAPMAN, R. F. 1982. The insects structure and function, 3rd ed. Harvard Univ. Press, 
Cambridge, Mass. 919 pp. 

CLAUSEN, C. P. 1940. Entomophagous insects. McGraw-Hill Book Co., New York. 688 








pp. 

Common, I. F. B. 1970. Lepidoptera in C.S.I.R.O. The insects of Australia. Melbourne 
Univ. Press, Carlton, Victoria. 1029 pp. 

1975. Evolution and classification of the Lepidoptera. Ann. Rev. Entomol. 20: 
183-203. 

COTTRELL, C. B. 1984. Aphytophagy in butterflies: Its relationship to myrmecophily. 
Zool. J. Linn. Soc. 79:1—57. 

Downes, J. A. 1978. Lepidoptera feeding at puddle-margins, dung and carrion. J. 
Lepid. Soc. 27:89-99. 

FEINSINGER, P. 1983. Coevolution and pollination. In Futuyma, D. J. and M. Slatkin 
eds. Coevolution. Sinauer Assoc. Inc. Sunderland, Mass. 555 pp. 

FARQUHARSON, C. O. 1922. Five years’ observations (1914-1918) on the bionomics of 
southern Nigerian insects, chiefly directed to the investigation of Lycaenid life his- 
tories and to the relation of Lycaenidae, Diptera and other insects to ants. Trans. 
Roy. Entomol. Soc. London 69:319-448. 

GILBERT, L. 1976. Adult resources in butterflies: African lycaenid Megalopalpus feeds 
on larval nectary. Biotropica 8:282-283. 

GRANT, V. & K. A. GRANT. 1965. Flower pollination in the phlox family. Columbia 
Univ. Press, New York. 180 pp. 

HAGEN, K. S. 1958. Honeydew as an adult fruit fly diet affecting reproduction. Proc. 
Int. Cong. Entomol. 10:25-30. 

1976. Role of nutrition in insect management. Proc. Tall Timb. Conf. Ecol. 
Anim. Contr. Habitat Mgmt. 4:221-261. 

HAGEN, K. S., P. GREANY, E. F. SAWALL & R. L. TASSAN. 1976. Tryptophan in artificial 
honeydews as a source of an attractant for adult Chrysopa carnea. Environ. Entomol. 
5:458-468. 

HEINRICH, B. 1979. Bumblebee economics. Harvard Univ. Press, Cambridge, Mass. 
245 pp. 

HENNING, S. F. 1983. Biological groups within the Lycaenidae. J. Entomol. Soc. S. Afr. 
46:65-85. 

HINTON, H. E. 1951. Myrmecophilous Lycaenidae and other Lepidoptera—a sum- 
mary. Proc. S. London Entomol. Nat. Hist. Soc. 1949-1950:111-175. 

JENSEN, W. A. & F. B. SALisBuRY. 1972. Botany: An ecological approach. Wadsworth 
Pub. Co., Belmont, Cal. 748 pp. 

LAMBORN, W. A. 1914. On the relationship between certain West African insects, 
especially ants, Lycaenidae and Homoptera. Trans. Roy. Entomol. Soc. London 61: 
436-524. 

NEUENSCHWANDER, P., K. S. HAGEN & R. F. SMITH. 1975. Predation on aphids in 
California’s alfalfa fields. Hilgardia 43:53-78. | 

OrsAk, L. J. 1977. The butterflies of Orange County, California. Misc. Publ. No. 3. 
Center for Pathobiology, Museum of Systematic Biology, Univ. of Cal., Irvine, Cal. 
349 pp. 

POWELL, J. A. 1980. Evolution of larval food preferences in Microlepidoptera. Ann. 
Rev. Entomol. 25:133-159. 

ROEPKE, W. 1918. Zur Myrmekophilie von Gerydus boisduvali Moore. Tijds. Entomol. 
61:1-16. 








VOLUME 39, NUMBER 4 AT 


STEBBINS, G. L. 1974. Flowering plants: Evolution above the species level. Harvard 
Univ. Press. 

STRONG, F. E. 1963. Studies on lipids in some homopterous insects. Hilgardia 34:43- 
61. 

VAN EMDEN, H. F. & K. S. HAGEN. 1976. Olfactory reactions of the green lacewing 
Chrysopa carnea to tryptophan and certain breakdown products. Environ. Entomol. 
0:469-473. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
39(4), 1985, 328 


NOTES ON THE PARASITISM OF ROTHSCHILDIA SP. PUPAE 
(SATURNIIDAE) IN GUANACASTE PROVINCE, COSTA RICA 


Tachinid flies (Diptera) comprise a major source of pupal mortality in Rothschildia 
spp. (Saturniidae) in El Salvador (Quezada, 1967, Ann. Entomol. Soc. Amer. 60:595- 
599). In spite of the broad geographical distribution of Rothschildia in Central America 
(Ferguson, 1972, The moths of America north of Mexico, Bombycoidea-Saturniidae, E. 
W. Classey, London, 275 pp.), little has been published on similar mortality agents on 
pupae for other localities. In this note I report some qualitative observations on pupal 
mortality in Rothschildia sp. from lowland Guanacaste Province, Costa Rica since (1) 
such data for this moth genus in Costa Rica are lacking, and (2) a hymenopterous species 
was discovered, providing a significant difference from the El Salvador studies (Quezada, 
op. cit.). 

Five intact cocoons of Rothschildia sp. were collected from one bush (1.5 m tall) in a 
roadside patch of deciduous forest about 5 km north of Bagaces (10°31'N, 85°15’W) along 
the Pan-American Highway on 2 March 1984. The cocoons were placed in a small “Zip- 
Loc” bag without close examination. About two months later (10 May) I noticed nu- 
merous newly emerged small wasps inside the bag. Upon closer examination I determined 
that a total of 67 wasps, all apparently the same species, emerged from two of the cocoons. 
All five cocoons were opened to determine the condition of the pupae. In one of the two 
cocoons from which wasps emerged, the pupa appeared mummified but with numerous, 
small round holes, apparently the exit sites of the wasps. The second parasitized pupa 
had no such holes but was broken open in the abdominal region. A single dead wasp was 
found at the bottom of the pupal cavity in each of these cocoons. Curiously and yet-to- 
be explained, a third cocoon was completely devoid of a pupa, pupal or larval exuvium, 
but had dried mud “caked”’ to the bottom of the pupal cavity. This cocoon also had a 
small round hole near the top (but not the emergence valve for the moth) tightly plugged 
with mud. This hole was about twice the diameter of the wasp emergence holes in the 
pupal cuticle found in one of the cocoons. A fourth cocoon contained a dead, mummified 
pupa, one dead wasp, and when broken apart, appeared to contain many mold spores. 
The fifth cocoon contained a hardened, mummified, but otherwise intact, dead pupa. 

The wasps were determined to be Spilochalcis sp. (Hymenoptera: Chalcididae). 

Although pupal parasitism in Rothschildia spp. in El Salvador is attributable primarily 
to tachinids, a low percentage of parasitism by an ichneumonid was also observed (Que- 
zada, op. cit.). Quezada does not mention chalcids as being a pupal parasite of these silk 
moths in El Salvador. While the cocoons in my study were clearly Rothschildia, it was 
not possible to confirm a species determination since no viable adults were obtained. 
Although my sample size is terribly small, it is also interesting to note that these pupae 
had been collected during the pronounced dry season of the region, but the adult parasites 
did not appear until the end of this period. 

I thank Susan S. Borkin for assistance and Dr. E. E. Grissell, Systematic Entomology 
Laboratory, U.S. Department of Agriculture, for determining the wasps. The wasps are 
deposited in the collections of the Milwaukee Public Museum. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


VOLUME 39, NUMBER 4 329 


Journal of the Lepidopterists’ Society 
89(4), 1985, 329-330 


AN ABERRATION OF ICARICIA ACMON LUTZI (LYCAENIDAE) 


Responding to the suggestion of F. Martin Brown, Colorado Springs, Colorado, I pres- 
ent a photograph of an aberrant male Icaricia acmon lutzi (dos Passos). This butterfly, 





Fic. 1. Aberrant male of Icaricia acmon lutzi, Custer Co., Idaho: Above—top view; 


below—bottom view. 


330 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the only one sighted to date, was captured while it was visiting cold campfire coals on 
14 July 1983, in Stanley Basin, Custer County, Idaho. Mr. Brown conveyed to me that 
he was unaware of any record of this aberrant form of lutzi. 


K. D. CANNON, 2420 Gekeler L., Boise, Idaho 83706. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 330-333 


NATURAL HISTORY NOTES FOR AELLOPOS CECULUS (CRAMER) 
(SPHINGIDAE) IN NORTHEASTERN COSTA RICA 


With the exception of one early study (Moss, 1920, Novit. Zool. 27:333—424) and one 
recent one (Haber & Frankie, 1983, In Janzen, D. H. (ed.), Costa Rican natural history, 
The Univ. of Chicago Press, Chicago, 816 pp.), little information has been published 
regarding the life cycle and associated natural history for Neotropical sphingids of the 
genus Aellopos (formerly Sesia). In this note I report additional information on the early 
stages and life cycle of A. ceculus (Cramer) (Fig. 1) at one locality in northeastern Costa 
Rica, including observations on oviposition, larval food plant, and caterpillar behavior. 
Previously, a description of the fifth instar larva and a larval food plant record had been 
reported by Moss (op. cit.) in Brazil. 

The locality is “Finca La Tigra,” near La Virgen (10°23'N, 84°07'W; 220 m eley.), 
Heredia Province and Sarapiqui District, Costa Rica. Information on this sphingid was 
generated by observing caterpillars in captivity and one instance of repeated oviposition 
attempts by one female in the wild. One fourth instar larva was reared in February 1984 
to adulthood, and a second individual was reared from egg to adult in August-September 
1984. Rearing was done by confining a caterpillar in a large, tightly closed, clear plastic 
bag containing fresh cuttings of the food plant. 

On 4 August 1984 and 1600 h, a female A. ceculus alighted a total of five times on 
the very long (approx. 1.0 m) meristem of the rubiaceous vine-like shrub Sabicea billosa 
R. & S. Immediately prior to this time, I observed the same moth meander through 
dense pockets of secondary-growth vines on the opposite side of the roadcut from this 
individual of S. billosa. The moth fluttered and hovered in this vine patch for several 
minutes before darting across the gravelly dirt road to oviposit on S. billosa. Although 
the moth momentarily alighted at several places on the long meristem, including un- 
furling leaflets, close examination of the vine following the departure of the moth re- 
vealed only a single egg carefully positioned on the dorsal surface of a tiny leaflet near 
the very tip of the meristemal growth (Fig. 2). Even though freshly opened flowers were 
present on the older portions of this vine and on adjacent individuals of S. billosa, A. 
ceculus did not pause to feed. Careful examination of both meristems and older leaves 
and flowers on three S. billosa vines in the same area revealed no additional eggs or 
sphingid caterpillars. All three vines possessed very clear evidence of recent “explosive” 
growth of meristems, easily recognizable by the reddish tinge of these tissues. 

The white 1.1 mm dia. spherical egg (Fig. 2) bears no external ridges or other sculp- 
turing and hatches in five days. The first instar larva immediately devours the egg shell; 
the larva is 6 mm long x 1.2 mm thick, and pale, translucent green with a 1.1 mm long 
terminal black caudal “horn.” About four days later, the caterpillar molts to the second 
instar; it is about 14 mm long at this time. Although very similar in overall appearance 
to the first instar, the caterpillar’s body cuticle assumes a reflective luster and with faint 
evidence of a medial, dorsal pink band running the length (Fig. 2). Throughout all instars, 
the head capsule remains pale green in color but with a stripe pattern becoming evident 
by the third instar larval stage. The trunk region of both the first and second instars is 
dark green and covered sparsely with fine setae. The caudal horn in both instars stands 


VOLUME 39, NUMBER 4 ool 





Fic. 1. Adult female Aellopos ceculus (Cramer), dorsal view, reared from the egg 
stage in this study. This specimen is deposited, along with the pupal shell, in the collec- 
tions of the Milwaukee Public Museum. 


almost perpendicular to the main axis of the body. The second instar lasts about five 
days and grows to 25 mm long. 

The third instar larva (Fig. 2) assumes the basic color pattern and overall appearance 
which is retained until pupation. The third instar attains a maximum body length of 31 
mm in about six days. The caudal horn is 5 mm long, deflected posteriorly, and reddish. 
The spiracles are also ringed in red. The broad dorso-medial band running lengthwise is 
faintly edged in white. This composite band begins on the second thoracic segment but 
becomes very pronounced on the abdominal segments. Laterally the trunk region is 
marked with a series of seven composite, oblique bands, reddish anteriorly and white 
posteriorly. These bands appear “white” even though they are actually composites of 
two colors (Fig. 2). Laterally, each of these composite oblique bands crosses three adjacent 
body segments and fuses into the dorso-medial line on the third segment (i.e., posterior- 
most) in such a triplet. Adjacent oblique bands overlap considerably in the segments 
bearing them (Fig. 2). All abdominal segments bear multiple vertical rows of pronounced 
white studs, readily seen with a 10x hand lens; these markings are also present on the 
thoracic segments, but they are less pronounced. The fourth and fifth instars are virtually 
identical in color patterns to the third instar. By the time of pupation, the caterpillar is 
51 mm long; the overall duration of the caterpillar stage is 28 days. 

A greenish prepupal stage lasts about three days, and the dark brown pupa measures 
31 mm long x 9 mm laterally through the wing pad (thoracic) area. In the rearing study, 
the pupa was formed in the folds of dampened paper towels at the bottom of the plastic 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Larval food plant, oviposition, and early stages for Aellopos ceculus at “Finca 
La Tigra,” near La Virgen, Sarapiqui District, Costa Rica. Top two photographs: the 
larval food plant, Sabicea billosa R. & S. (Rubiaceae) showing the long meristem and 
terminal leaflet used as an oviposition site (left), and position of the egg on the dorsal 
surface of an unfurling meristemal leaflet (right). Left column below these photographs: 
close-up view of the egg, and second instar larva (dorsal view) and resting on ventral 
midrib of meristemal leaf; third instar larva, lateral view, showing pattern of markings 
and studded rugosity of the cuticle (right); also resting on a midrib of a food plant leaf. 





VOLUME 39, NUMBER 4 333 


bag containing the food plant. When handled, the pupa quickly responds with violent 
movements of the abdominal segments. The pupa stage lasted 22 days for my sample of 
N = 1 individual reared from egg to adult. A pupa obtained in February 1984 from a 
caterpillar discovered in the fourth instar lasted 17 days. The individual A. ceculus 
obtained from the complete rearing (Fig. 1) was a female. This individual has a wing- 
spread of 39 mm and a body length of 25 mm. A detailed description of the adult for 
this species (under Sesia) is given in Seitz (1924, Macrolepidoptera of the World, vol. 6, 
A. Kernan, Stuttgart, 501 pp.). 

Following the devouring of the egg shell, the first instar larva positions itself along the 
midrib of the ventral surface of a reddish meristemal leaf. The caterpillar throughout 
all five instars perches on the midrib of a leaf and appears very cryptic in this manner. 
Until the latter half of the third instar, the caterpillar feeds exclusively on reddish, soft 
meristemal leaf tissues and not on the greener, mature leaves of S. billosa. Feeding 
appears to occur in both day and night. 

This sphingid is generously distributed geographically from Mexico to southern Brazil 
(Seitz, op. cit.). Moss (op. cit.) reported two different Rubiaceae as larval food plants in 
Brazil, including Sabicea. Moss reported that different color morphs of the caterpillar 
are associated with the two different food plants, and the description for the Sabicea- 
associated form matches that reported here for the Costa Rican population. One inter- 
esting difference, however, is that Moss reported Sabicea-associated caterpillars to possess 
chestnut-red lateral stripes, while my individual clearly had more whitish stripes. Haber 
and Frankie (op. cit.) also note that the caterpillars of A. titans Cramer are dimorphic 
in color. Larval descriptions and food plants for other species of Aellopos are summarized 
in Hodges (1971, The moths of America north of Mexico, Fasc. 21, Classey, London, 158 
pp.). But, Hodges (op. cit.) also reports that descriptions of early stages and associated 
natural history data are poorly known for some species. In Costa Rica, A. ceculus has 
previously been reported for a different, lower-elevation locality in the Sarapiqui District, 
“Finca La Selva,” about 25 km from “Finca La Tigra,” and it is one of five species of 
the genus known to cccur in Costa Rica (Haber, 1988, In Janzen, D. H., op. cit.). Inter- 
estingly, A. ceculus is the only member of the genus reported from the Sarapiqui District 
(Haber, op. cit.). 

Oviposition behavior in the late afternoon hours may be part of a more general diurnal 
pattern of adult activity in this sphingid in which activities are concentrated near dusk 
(e.g., Haber and Frankie, op. cit.). Although Aellopos adults may feed on the flowers of 
their larval food plants (Haber and Frankie, op. cit.), this behavior was not observed in 
my very brief study, even though flowers were present on the S. billosa vines examined. 
And while most sphingids, including Aellopos, presumably place their eggs on the un- 
dersides of leaves on the larval food plant (e.g., Haber & Frankie, op. cit.), this was not 
the case for A. ceculus in the present study, albeit a very limited sample. Aellopos ceculus 
gravid females in search of egg-placement sites, as well as other sphingids in the Neo- 
tropical Region, may opportunistically exploit meristemal tissues of larval food plants 
proliferating at certain times of the year. These tissues may serve as highly suitable 
oviposition sites in that newly hatched caterpillars are placed in close spatial proximity 
to soft, digestible plant tissues under these conditions. Since there is little evidence in- 
dicating that Neotropical sphingid adults or their caterpillars are unpalatable as a con- 
sequence of larval food plant selectivity (Haber & Frankie, op. cit.), observed oviposition 
preference by A. ceculus, and perhaps other sphingids, for seasonally available meriste- 
mal tissues may indicate a facultative form of natural selection permitting these herbiv- 
orous organisms to exploit the most energetically cost-efficient food plant tissues. 

Susan Borkin and Joan Jass discovered the A. ceculus caterpillar in February 1984, 
and they shared their observations with me. Luis Diego Gomez of the Herbarium, Na- 
tional Museum of Costa Rica, identified the larval food plant based upon the examination 
of extensive vegetative material as well as flowers. Mel Scherbarth photographed the 
adult moth illustrated in this note. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 582838. 


334 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Journal of the Lepidopterists’ Society 
89(4), 1985, 334 


MISDIRECTED MONARCH MATING BEHAVIOR 
(DANAIDAE: DANAUS PLEXIPPUS) OR NOBLESSE OBLIGE? 


T. E. Pliske (1975, Ann. Entomol. Soc. Amer. 68:143-151) has described the aggres- 
siveness of the male Danaus plexippus (Linn.) as it employs its “take-down’” maneuver 
to drive the female to the ground during its courtship routine. J. W. Tilden (1979 (81), 
J. Res. Lepid. 18:2) has depicted the male’s sometimes faulty discrimination resulting in 
male attempting to mate with male. That this indiscriminate activity can become even 
more misdirected is evidenced by the following observation. 

In April 1982, while photographing butterflies nectaring at a Pittosporum tree on 
Ossabaw Island, Chatham County, Georgia, I noted two D. plexippus, the first a worn 
and decrepit female and the other a fairly fresh and active male, neither of which showed 
any interest in the other. 

As I watched, the cruising male suddenly stooped like a falcon, struck a nectaring 
Vanessa virginiensis (Drury), sex undetermined, from its blossom, and pinned it to the 
pavement below (Fig. 1). In the brief moment available for photographing the event, I 
did not observe any actual attempt to copulate. The virginiensis then struggled free, and 
both butterflies flew off. 

One should perhaps resist the temptation to anthropomorphize regarding the monarch 
and the painted lady. 


DAVE WINTER, 257 Common St., Dedham, Massachusetts 02026. 





Fic. 1. V. virginiensis, grasped by the legs of a male D. plexippus and pinned to the 
ground. 


VOLUME 39, NUMBER 4 ODD 


Journal of the Lepidopterists’ Society 
89(4), 1985, 335 


A NEW FOOD PLANT RECORD FOR ATALOPEDES CAMPESTRIS 
(BOISDUVAL) (HESPERIIDAE) 


Atalopedes campestris (Boisduval) is a common skipper found in the new world from 
Canada to Ecuador and northern Brazil (Evans, 1955, A catalogue of the American 
Hesperiidae, the British Museum, London). Host plant records for the larval stages of A. 
campestris include several grass species: 1) Bermudagrass, Cynodon dactylon (L.) Pers. 
(Klots, 1951, Field guide to the butterflies, Houghton-Mifflin, Co.; Warren & Roberts, 
1956, J. Kans. Entomol. Soc. 29:189-41; Harris, 1972, Butterflies of Georgia, Univ. Okla- 
homa Press); 2) St. Augustinegrass, Stenotaphrum secundatum (Walter) Kuntze (Howe, 
1975, Butterflies of North America, Doubleday & Company, Inc.); 3) large crabgrass, 
Digitaria sanguinalis (L.) Scop.; and 4) saltgrass, Distichlis spicata (L.) Greene (Tietz, 
1972, An index to the described life histories, early stages, and hosts of Macrolepidoptera 
of the continental United States and Canada, Allyn Mus. Entomol., Sarasota, FL). 

Several “tent’”’ structures typical to grass and sedge feeding Hesperiinae were observed 
on biotypes of Cogongrass, Imperata cylindrica (L.) Beauv. on 28 September 1984 at 
Stoneville, MS. Two larvae and one pupa were found, and from these, two male and one 
female A. campestris adults emerged on 5 and 10 October. Additional larvae and “tents’’ 
were observed on I. cylindrica biotypes collected from Alabama, Mississippi (Patterson, 
1980, Proc. So. Weed Sci. Soc. 33:251) and Iraq (Al-Juboory & Hassaway, 1980, Weed 
Sci. 28:324—26). 

These observations not only establish a new host plant record for A. campestris but 
indicate that this skipper should be evaluated for its potential as a biological control agent 
against I. cylindrica. Biological controls are certainly needed for this weedy native of 
Indo-Malaysia. It is an aggressive, rhizomatous perennial weed, ranking as the world’s 
seventh worst weed (Holm et al., 1977, The World’s worst weeds, The University Press 
of Hawaii). Since its introduction between 1910 and 1920 (Patterson, 1980, Weed Sci. 
28:735-740), it has become a pernicious weed of non-cultivated areas in the southeastern 
United States. 


CHARLES T. BRYSON, USDA-ARS, Southern Weed Science Laboratory, Stoneville, 
Mississippi 38776. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 385-337 


SPECIMENS OF CALLOPHRYS RUBI L. (LYCAENIDAE) FROM FIJI— 
TRANSPLANTED COLONY OR ONE-TIME OCCURRENCE? 


While studying specimens of worldwide Callophrys-related taxa in the British Museum 
(Natural History) in 1983, I located two specimens of C. rubi L. in unincorporated 
material of the Adams Bequest, which bore labels indicating capture in Fiji in 1904. 
Given the oddity of these data on specimens of a butterfly generally distributed from 
the British Isles eastward through Soviet Asia (Higgins & Riley, 1970, A field guide to 
the butterflies of Britain and Europe, Houghton-Mifflin Co., Boston; Johnson, 1986, A 
revision of the Callophryina of the world with phylogenetic and biogeographic analyses, 
Bull. Am. Mus. Nat. Hist., in press), the specimens were photographed (Fig. 1). Robinson 
(1975, Macrolepidoptera of Fiji and Rotuma, Classey, London) does not list C. rubi from 


336 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


eG ee 


ae: Cubi rx-11-04 












: Adams Bequest. 
B.iM. 1942-399, 


Fic. 1. Photograph of two male specimens of C. rubi and data recording their capture 
in Fiji. Specimens are in British Museum (Natural History). 


Fiji. He records 45 species of butterflies for the island, of which four are cited as endemic. 
It was, therefore, important to ascertain the possible validity of the above-mentioned 
specimens and their associated data. Two factors are relevant to this consideration—the 
overall veracity of data in Adams Bequest material and the availability of suitable larval 
foodplants in Fiji to support C. rubi. Regarding the former, I have examined Adams 
Bequest material from some 18 genera of Lycaenidae in the British Museum. Although 
some data are limited to only regional or country citation, I have never found an instance 
suggesting erroneous data. On the contrary, when Adams Bequest material has provided 
examples of species poorly represented in international collections, such material has 
always had data compatible with the known distributions of such species. Further, the 
C. rubi specimens noted above were found with other Fiji material, including the ly- 
caenids Zizina otis mangoensis (Butler) and Strymon bazochii gundlachianus (Bates), 
both listed by Robinson from Fiji. Regarding the question of suitable larval foodplant 
availability, Dr. Herbert Wagner (University of Michigan, Ann Arbor) has informed me 
that of the foodplants of C. rubi listed by Tutt (1907-1908, British Lepidoptera, Swan 
Sonnenschein & Company, London [p. 109]) the following are known to have been 
transplanted to Fiji and, consequently, occur there in varying distributions: Rubus idaeus 
(Richter), R. frangula (Glitz), Rumex spp., Medicago lupulina Linnaeus, Lotus cornic- 
ulatus Linnaeus, Trifolium spp., Genista tinctoria Linnaeus, Cytisus spinosus (Lin- 
naeus), and Amygdalus spp. The above categories of taxa represent some 25% of the 
larval foodplants listed by Tutt. Considering the above, it seems reasonable to accept the 
two British Museum specimens of C. rubi from Fiji as probably valid records. They have 
been curated by me into the overall collection of C. rubi at that museum with a special 
label citing this present note. It remains to be resolved whether this occurrence represents 
a possible transplanted colony of C. rubi in Fiji or simply a one-time occurrence due to 
accidental transplantation. Robinson (loc. cit.) cites human factors as having massive 
influence upon the fauna and flora of Fiji. He also records some butterflies of Fiji as 
known only from original types or (as in the case of Nacaduba dyopa (Herrich-Schaffer)) 
as having representation by a large series from one time with few, if any, subsequent 
captures recorded. Considering the above and the availability of Robinson’s general 


VOLUME 39, NUMBER 4 oor 


faunal work, the publication of these data concerning C. rubi specimens from Fiji has 
seemed advisable. 


KURT JOHNSON, Dept. of Entomology, American Museum of Natural History, Cen- 
tral Park West at 79th St., New York, New York 10024. 


Journal of the Lepidopterists’ Society 
39(4), 1985, 337-338 


ECOLOGICAL OBSERVATIONS ON APODEMIA PHYCIODOIDES 
BARNES & BENJAMIN (RIODINIDAE) 


In their paper on the rediscovery of Apodemia phyciodoides Barnes & Benjamin, 
Holland and Forbes (1981, J. Lepid. Soc. 35:226—-232) indicated that the ecological as- 
sociations of phyciodoides were imperfectly known and required further study. In late 
July 1984, I was joined by three other members of the Arizona Entomological Society 
on a trip to southeastern Sonora, Mexico. This trip was part of an on-going study of the 
Lepidoptera of Sonora by several members of the Arizona group. Observations made 
during this trip on phyciodoides may further clarify its habitat preferences and relation- 
ships with other riodinids. 

Our group consisted of Jim Brock, John Palting, Steve Prchal and myself. The eight 
day collecting trip was spent along Highway 16, southeast of Hermosillo, terminating at 
Yecora, near the Chihuahua state line. The collecting area covered was primarily in the 
Sierra Madre Occidental and its outer foothills and was about 125 airmiles south of the 
area collected by Holland and Forbes. Four biotic communities (or life zones) were 
sampled, from San Jose de Pimas to Yecora. Using terminology from Brown (ed., 1982, 
Desert plants 4:1-342, Biotic communities of the American southwest—United States 
and Mexico), these communities were: Sinaloan Thornscrub, Sinaloan Deciduous Forest, 
Madrean Evergreen Woodland, and Petran Montane Conifer Forest. A. phyciodoides 
was found to be relatively common along the dirt road between Santa Rosa and Yecora, 
from four to 10 miles east of Santa Rosa. All of these sites fall in the Madrean evergreen 
woodland community, a Quercus-Juniperus-Pinus habitat. The lowest collecting site, 
four miles east of Santa Rosa, is near the transition into the Sinaloan deciduous forest, 
which is indicated by a Ficus-Ceiba-Celtis habitat. The upper collecting site, 10 miles 
east of Santa Rosa, is near the plateau region of the Petran montane conifer forest, 
dominated by Pinus species. The Quercus dominated habitat in which phyciodoides was 
found consisted of a rugged canyon-ridge (barranca) geography. Despite fairly extensive 
collecting, phyciodoides was not observed in either of the two adjoining biotic commu- 
nities. Other “indicator” butterfly species that were most prevalent in the Madrean 
evergreen woodland were: Thessalia theona ssp. (Menetries), Thessalia cyneas (Godman 
& Salvin), an unknown Piruna species, and Cyllopsis pyracmon nabokovi L. Miller. The 
presence of phyciodoides and cyneas together, both of which previously were found in 
the Chiricahua Mountains of Arizona and both of which have not been found there 
recently, is intriguing. 

On 29 July 1984, while travelling up toward Yecora, both sexes of phyciodoides were 
observed in mid-afternoon at wet places along the dirt road. On 30 July, Brock and I 
hiked about four miles down the road in late morning, starting from the upper collecting 
site. The entire length of the hike was in the Quercus woodland habitat. Males of phy- 
ciodoides were observed patrolling along, and landing in, the dirt road. Both sexes were 
also observed at moisture and nectar. A total of about 25 specimens were collected in 
the two days. Extensive collecting in the Sinaloan deciduous forest (one to three miles 
east of Santa Rosa) on 29 July yielded no specimens of phyciodoides. The conifer forest 


338 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


plateau above the Quercus zone was also sampled without any observations of phycio- 
doides. The results of this sampling would indicate that the species is associated with the 
Quercus dominated Madrean evergreen woodland biotic community. According to Brown 
(1982, ibid.), this community also dominates much of the region collected by Holland 
and Forbes, as well as the Chiricahua Mts. of Cochise Co., Arizona—the type locality 
for phyciodoides. 

Holland and Forbes also expressed interest in the relationship between phyciodoides 
and other riodinids, especially Apodemia palmeri (Edwards) and A. hepburni (Godman 
& Salvin). While our limited observations cannot answer the questions in this subject, 
they may provide a better understanding of the Riodinidae associations by biotic com- 
munity. In addition, a potential pattern becomes apparent in the associations between 
phyciodoides, palmeri, and hepburni. 

No riodinid species were observed in the sampling of the Petran montane conifer 
forest. Within the Madrean evergreen woodland, we found phyciodoides, Apodemia 
hypoglauca (Godman & Salvin), Emesis ares (Edwards), Baeotis zonata (Felder), Lasaia 
maria Clench, and Calephelis arizonensis McAlpine. Only phyciodoides and arizonensis 
were common in this area, the other species were represented by less than half a dozen 
specimens each. In the Sinaloan deciduous forest, and lower into its transition with the 
Sinaloan thornscrub (16 miles east of Tecoripa to three miles east of Santa Rosa), six 
species were recorded: hepburni, hypoglauca, arizonensis, maria, Emesis poeas, and 
zonata. In this zone, only maria and hepburni were common. A. hypoglauca was not 
observed here on the late July 1984 trip but was taken on 26 August 1984 by Doug 
Mullins within the same zone (near Tepoca, highway 16). A stop along the river at San 
Jose de Pimas in the lower Sinaloan thornscrub (transitioning to Sonoran desert-scrub) 
resulted in three different species: palmeri, Apodemia mormo mejicanus (Behr), and 
Calephelis nemesis (Edwards). A. palmeri was common while the other species were 
present in lower numbers. 

In summation, these observations do not answer the problems presented by Holland 
and Forbes, but they do provide further clarification of the biotic associations of phycio- 
doides and its relationship with palmeri and hepburni. In southeastern Sonora, phycio- 
doides appears to be closely associated with the Quercus dominated Madrean evergreen 
woodland, a habitat which dominates middle and upper mountain regions into south- 
eastern Arizona. Categorizing riodinid species by biotic community in this region gives 
the perception that the three Apodemia species mentioned above are each associated 
with different habitats, and perhaps “replace” each other as the biotic communities are 
transversed. 


MICHAEL J. SMITH, 5407 Orinda Ave., Las Vegas, Nevada 89120. 


Journal of the Lepidopterists’ Society 
39(4), 1985, 338-339 


POPULATION OUTBREAK OF PANDORA MOTHS 
(COLORADIA PANDORA BLAKE) IN THE 
MAMMOTH LAKES AREA, CALIFORNIA 


Pandora moths (Coloradia pandora Blake), which are fairly widespread over the north- 
ern pine forests of the west, periodically exhibit an unusual increase in population as 
described by Brown (1984, J. Lepid. Soc. 38(1):65) and Ferguson (1971, Moths of America 
north of Mexico, Fasc. 20.2a, E. W. Classey Ltd., London). During a field trip in 1982, 
shortly after the described outbreak of adults on the Kiabab Plateau of Arizona, such an 
outbreak was witnessed in the Mammoth Lakes area (el. 7000 ft.) of California. On the 


VOLUME 39, NUMBER 4 339 


night of 30 August 1982, many hundreds of adults were observed flying and at rest on 
a motel in the town of Mammoth. Activity began at about 2000 h and continued for at 
least several hours. 

At a rest stop located five miles north of Mammoth on route 395 at the same approx- 
imate altitude, several thousand adults were seen the next day (31 August 1982). On the 
ground of the north side of the rest stop building were many hundreds of bodies and 
fragments of bodies, indicating probable predation. This evidence consisted of disasso- 
ciated heads and wings covering a large area. 

Activity at the motel resumed the night of the 31st, and several females were captured. 
Each of these laid up to a hundred blue-green spherical eggs, which were not kept 
through hatching. 

The area around Mammoth is covered almost exclusively with lodgepole pine (Pinus 
contorta), and this forest, one of the largest in California, extends past the rest stop 
mentioned above. 

As a collector’s note, the rest stop described above has proven to be an excellent 
collecting spot, when open, which depends on enough water being available to make it 
usable. Many specimens can be taken there, including large Saturnidae, as the building 
is lit at night. Also, less than two miles north of the Rest Stop, route 395 crosses Deadman’s 
Creek, an excellent collection area for butterflies. 


KELLY RICHERS, 5913 Bel Aire Way, Bakersfield, California 93309. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 8339-340 


PUDDLING BY SINGLE MALE AND FEMALE TIGER SWALLOWTAILS, 
PAPILIO GLAUCUS L. (PAPILIONIDAE) 


The eastern tiger swallowtail, Papilio glaucus L., is noted for puddling in large groups 
on damp soil. These conspicuous aggregations are apparently all male; no female has 
ever been reported in them. This agrees with the general case in the Lepidoptera. In 
both butterflies and moths, puddling is a far more common behavior in males than in 
females (Downes, 1978, J. Lepid. Soc. 27(2):89-99; Adler, 1982, J. Lepid. Soc. 36(3):161- 
178). 

Puddling is apparently associated with the acquisition of sodium ions and amino acids 
from the substrate (Arms et al., 1974, Science 185:372-374). Adler and Pearson (1982, 
Can. J. Zool. 60:322-325) have shown that the sodium budgets of males and females of 
the cabbage butterfly, Pieris rapae L., are significantly different, with males having a 
higher need for sodium than females. This greater need for sodium by males may reflect 
the more active role of the male in reproduction, both in terms of greater flight activity 
(Downes, op. cit.) and in the production of nutrient rich spermatophores (Adler & Pear- 
son, op. cit.). This in turn may explain the preponderance of males at puddling aggre- 
gations. 

We have eight observations of fresh male P. glaucus puddling singly over the course 
of several summers near Ithaca, Tompkins Co., N.Y. and near Cooperstown, Otsego Co., 
N.Y. These may represent cases where the individual is the first to find an area of rich 
resources and thus may form the core of a puddling aggregation later on. Males in this 
species are attracted to conspecific decoys (Arms et al., op. cit.). This may be a conse- 
quence of their mate-locating behavior, which apparently involves searching for mates 
at a wide variety of sites (Berger, pers. comm.). Patrolling males may key onto a puddling 
individual in the hopes that it is a female and remain at the puddling site if it is rich in 


340 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


needed nutrients. Alternatively, single puddling males may be at sites with lower con- 
centrations of the needed resources. 

We have observed five cases of puddling by female P. glaucus at a study site near 
Cooperstown, Otsego Co., N.Y. In all five cases, the females were puddling singly. The 
first observation was at 1050 h on 22 June 1983 (day 16 of the brood). A fresh female 
was captured while puddling on damp soil in a vegetable garden, where she had been 
settled for about two minutes. 

Three observations all occurred on 18 June 1984 (day 10 of the brood). At 1045 h, a 
fairly worn female was disturbed while puddling on damp soil at the edge of a road. 
She flew to the end of a nearby cornfield where she puddled in two different locations 
for a total duration of about five minutes. This female was subsequently captured while 
nectar-feeding. At 1230 h, a fresh female flew slowly along a different road edge. She 
landed once, probed at the soil, then continued down the road. Finally, at 1620 h, a 
slightly worn female was seen taking off and landing several times along the road edge, 
probing the soil at least once. 

The fifth observation was at 1627 h on 2 June 1985 (day 17 of the brood). A very worn 
female was observed taking off and landing at several different spots on the soil of the 
vegetable garden and was subsequently captured after she had been puddling for about 
three minutes. 

Papilio glaucus is a highly vagile, wide-ranging species. Both sexes show very low 
recapture rates in mark-recapture studies (Lederhouse, 1982, Ecol. Entomol. 7:379-383). 
Females of this species may well have greater relative nutrient requirements than females 
of more sedentary species. Puddling females may represent those cases where their 
requirements cannot be met from larval feeding, nectar, or the contributions of a male’s 
spermatophore. 

However, we have observed both males and females puddling singly in two related 
species, the black swallowtail, Papilio polyxenes F., and the zebra swallowtail, Eurytides 
marcellus (Cramer). The black swallowtail is not a wide-ranging species and differs 
considerably from the tiger swallowtail in its habitat preference and reproductive strat- 
egy (Lederhouse, 1983, Oecologia 59:307-311). That males puddle singly in this species 
may again be influenced by their territorial mating system, which involves male defense 
of lek sites (Lederhouse, 1982, Behav. Ecol. Sociobiol. 10:109-118). The observations of 
females puddling in these species may suggest that puddling in female Lepidoptera is 
more common than is widely believed. 

One possible reason why female Lepidoptera are not often seen puddling in groups | 
may be to avoid harassment by males at these sites. We have often observed the inves- 
tigation of and attempted copulation with puddling individuals by new arrivals at ag- 
gregations of puddling P. glaucus males. A female in this situation would have to com- 
promise between efficient puddling and exercising her reproductive choice. 


THERESA A. BERGER AND ROBERT C. LEDERHOUSE, Department of Zoology and 
Physiology, Rutgers University, Newark, New Jersey 07102. 


Journal of the Lepidopterists’ Society 
39(4), 1985, 340-341 


FLOWER VISITATION RECORDS FOR SNOUT 
BUTTERFLIES (LIBYTHEIDAE) 


In the course of a general survey of libytheid butterflies (Shields, Tokurana, in press), 
flower visitation records were noted for Libytheana bachmanii Kirtland (most) and Li- 
bythea celtis Fuessly, gleaned from published sources and correspondence. These records 
are arranged here according to the classification of A. Takhtajan (1969, Flowering plants: 


VOLUME 39, NUMBER 4 341 


Origin and dispersal, Smithsonian Institution Press, Washington, D.C., 310 pp.), from 
primitive to advanced: 


Clematis vitalba (Ranunculaceae) 
Boussingaultia leptostachya (Basselaceae) 
Eriogonum sp. (Polygonaceae) 

Erica cinerea (Ericaceae) 

Tilia sp. (Tiliaceae) 

Croton sp. (Euphorbiaceae) 

Rubus sp. (Rosaceae) 

Prunus caroliniana (Rosaceae) 
Eysenhardtia amorphoides (Leguminosae) 
Melilotus albus (Leguminosae) 
Philadelphus coronarius (Saxifragaceae) 
Cornus sp. (Cornaceae) 

Ligustrum vulgare (Oleaceae) 

Baccharis sarothroides (Compositae) 
Senecio douglasii (Compositae) 
Chrysothamnus sp. (Compositae) 
Cirsium sp. (Compositae) 

Mentha sp. (Labiatae) 

Sorghum sp. (Gramineae) 


It is instructive to compare this list with flower visitation records for Asterocampa by 
Neck (1983, J. Lepid. Soc. 37:269-274), another nymphalid genus that utilizes Celtis 
(Celtidaceae) for larval foodplants. The only overlap in nectar feeding for both was 
Leguminosae and Saxifragaceae. However, Asterocampa adults were also reported on 
the fruit of Rubus and Prunus (Rosaceae), two genera that appear in the flower visits 
for libytheids. These facts may take on phylogenetic significance, since Libythea celtis 
uses both Celtis australis L. and Prunus as larval foodplants (Vladimir B. Polacek, in 
litt.); and Rosaceae, Leguminosae, and Saxifragaceae are closely related. A preliminary 
survey of flower visitation records for butterflies appears in Shields (1972, Pan-Pac. Ento- 
mol. 48:189-203). 


OAKLEY SHIELDS, 4890 Old Highway, Mariposa, California 95338. 


Journal of the Lepidopterists’ Society 
89(4), 1985, 341-342 


MALE DETERMINED MATING DURATION IN BUTTERFLIES? 


When considering what factors influence the mating duration in butterflies, it is im- 
portant to know to what extent it can be influenced by each sex respectively. Sims (1979, 
Am. Midl. Nat. 102:36-50) suggested, in analogy with results by Leopold, Terranova and 
Swilley (1971, J. Exp. Zool. 176:353-360) on Musca domestica, that mating duration 
probably is controlled by the female. This may be true in the sense that the female can 
inform the male when she is ready to terminate the copulation. However, in butterflies 
it is more likely that the male ultimately determines mating duration. If there should 
exist a conflict between the male and the female about when to terminate the copulation, 
the construction of the male genitalia suggests that the male alone determines copula 
duration. This inference is supported by two incidental observations I have made. 

The first concerns a pair of Coenonympha pamphilus (Satyridae), where the female 
was killed during copulation. On 26 August 1982 in Timmernabben, Sweden, I released 


342 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a virgin female to a male. After mating for 1 h 89 min a crab spider (Thomisidae) 
attacked and grabbed the female. After 4 h 51 min I left them while they were still in 
copula but put a cage over them. When I returned half an hour later the copulation was 
over; the male was flying in the cage while the female was still held by the spider among 
the vegetation. 

The other observation concerns a pair of Pararge aegeria (Satyridae) kept in a cage 
in the laboratory, where the male for unknown reasons died during copulation. They 
were found in copula on 5 February 1985 at 1100 h. It was still dark in the cage, and 
the pair must have been mating since 1840 h the day before when the light was switched 
off automatically. At 1340 h the light was turned on. At 1700 h the female was found 
flying in the cage with the male hanging from the tip of her abdomen. Upon inspection 
he was found to be dead. On 8 February the female was still alive and attached to the 
dead male. On 15 February the female was found dead, still in copula with the male. 

Although not proven, it seems plausible that the copulation is ultimately terminated 
by the male. Further observations and experiments would be interesting. 


PER-OLOF WICKMAN, Department of Zoology, University of Stockholm, S-106 91 
Stockholm, Sweden. 


Journal of the Lepidopterists Society 
39(4), 1985, 342 


SATYRIUM AURETORUM AURETORUM (BOISDUVAL): A NEW 
SPECIES FOR OREGON (LYCAENIDAE) 


The senior author received a number of butterflies from the junior author among 
which were three female Satyrium auretorum auretorum (Boisduval) with the following 
data: OREGON: Lake Co.; 2 miles south of Lakeview, 1 June 1981, leg. Ray Albright. 
Dornfeld (1980, The butterflies of Oregon, Timber Press, Forest Grove, OR) does not 
report the species for the state. The species has been expected there; it was taken towards 
the border in Siskiyou Co., California (Klamath River, near I-5, fide S. O. Mattoon). This 
new location also brings the species to within 20 miles of the Nevada border, another 
state where it is unrecorded. 

We thank S. O. Mattoon for providing the northern California record. 


GEORGE T. AUSTIN, Nevada State Museum and Historical Society, 700 Twin Lakes 
Drive, Las Vegas, Nevada 89108 AND RAY ALBRIGHT, Rt. 1, Box 277, Dayton, Oregon 
97114. 


Journal of the Lepidopterisis’ Society 
39(4), 1985, 343-344 


INDEX TO VOLUME 39 


(New names in boldface) 


Aellopos ceculus, 330 Godfrey, G. L., 57 
Agathymus, 171 Hamadryas, 229 
Agrias, 266 Hammond, P. C., 156 
Amphion nessus, 53 Hawkeswood, T. J., 276 
Anisota virginiensis, 53 Heliconiidae, 95 
Apantesis parthenice, 59 Hesperiidae, 48, 62, 95, 215, 239, 284, 299, 
Apaturidae, 95, 284 335 
Apodemia phyciodoides, 337 Higgins, L., 145 
Arctiidae, 59, 239 Hodges, R. W., 189, 151 
Astraptes, 215 Hyalophora cecropia, 65 
Atalopedes campestris, 335 H. columbia, 65 
Austin, G. T., 95, 342 Icarica acmon lutzi, 329 
Automeris iris hesselorum, 168 Incisalia henrici, 62 

A. randa, 163 Jalmenus daemeli, 276 
Battus philenor, 228 Johnson, J. B., 321 
Berger, T. A., 339 Johnnson, K., 119, 335 
Blanchard, A., 1 Kellogg, T. A., 268 
Book review, 51, 237 Knudson, E. C., 1 
Borkin, S. S., 229 Koerber, T. W., 26 
Brassolidae, 33, 225 Lederhouse, R. C., 339 
Brower, A. E., 280 Libytheidae, 95, 284, 340 
Brown, L. N., 196, 262 Lycaeides argyrognomon, 145 
Bryson, C. T., 335 Lycaenidae, 62, 95, 119, 145, 239, 276, 284, 
Butler, L., 177 329, 335, 342 
Calhoun, J. V., 284 Mather, B., 134 
Callophrys rubi, 335 Mather, K., 134 
Cannon, K. D., 329 May, P. G., 53 
Carmeuta laurelae, 262 McCorkle, D. V., 156 
Carter, M., 125 Megathymidae, 171 
Catocala, 280 Miller, L. D., 187 
Cock, M. J. W., 48 Mitoura millerorum, 119 
Coloradia pandora, 338 Moodna bisinuella, 9 
Cossidae, 1 Morphidae, 239 
Cubero, R., 33 Neck, R. W., 228 
Cymaenes finca, 48 Neunzig, H. H., 9 
Danaidae, 95, 239, 284, 334 Nielsen, M. C., 62 
Danaus plexippus, 334 Noctuidae, 1, 48, 57, 239, 280, 321 
Daterman, G. E., 26 Notodontidae, 1 
Davis, D. R., 235 Nymphalidae, 55, 95, 146, 239, 266, 284, 
Dicymolomia metalliferalis, 13 229 
Dowell, R. V., 237 Occidryas anicia bernadetta, 55 
Eichlin, T. D., 196, 262 Opsiphanes, 225 
Eisele, R. C., 238 O. quiteria quirinus, 38 
Eriocrania, 52 Papilio glaucus, 339 
Eriocraniidae, 52 P. polyxenes asterius, 125 
Eryphanis aesacus buboculis, 33 Papilionidae, 95, 125, 156, 228, 239, 284, 
Eucosma, 26 339 
Eurema nise, 238 Papilionoidea, 19 
Evans, D. L., 43 Parnassius clodius, 156 
Exoteleia anomala, 139 Pavulaan, H., 19 
Feeny, P., 125 Pedaliodes perperna, 187 
Gelechiidae, 151, 189 P. petronius kerrianna, 187 


Geometridae, 1, 145, 177, 239 P. petronius petronius, 187 


344 


Phigalia titea, 177 
Pieridae, 95, 238, 239, 268, 284 
Pieris rapae, 239 
P. sisymbrii, 268 
Powell, J. A., 26 
Pseudosphinx tetrio, 208 
Pygrus centaureae, 62 
Pyralidae, 1, 9, 13 
Ray, T. S., 266 
Rhyacionia, 26 
Richers, K., 338 
Rindge, F. W., 145 
Riodinidae, 95, 224, 337 
Robbins, R. K., 224 
Rothschildia, 328 
Santiago-Blay, J. A., 208 
Sartwell, C., 26 
Saturniidae, 53, 65, 85, 163, 328, 338 
Satyridae, 95, 187, 239, 284, 341 
Satyrium auretorum auretorum, 342 
Scelionidae, 59 
Sesiidae, 196, 262 
Shields, O., 340 
Smith, M. J., 337 
Snow, J. W., 196, 262 
Sower, L. L., 26 
Sphingicampa albolineata, 85 
S. montana, 85 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Sphingidae, 1, 53, 208, 330 
Spomer, S. M., 55 

Stafford, M. P., 321 
Stallings, D. B., 171 
Stallings, V. N. T., 171 
Stevens, R. E., 26 
Synanthedon dominicki, 196 
Telenomus, 59 

Tigridia acesta, 146 
Tildenia georgei, 151 
Tortricidae, 1, 26 

Turner, B. R., 171 

Turner, J. R., 171 

Turner, J. R. G., 201 
Tuskes, P. M., 85, 163 
Tuttle, J. P., 65 

Urbanus, 215 

Wagner, D., 13, 52 
Walker, T. J., 313 
Wasserman, F. E., 239 
Wickmann, P.-O., 341 
Williams, B. D., 53 
Winter, D., 334 

Wourms, M. K., 239 
Young, A. M., 146, 215, 225, 229, 328, 330 
Zygaenidae, 239 


Date of Issue (Vol. 39, No. 4): 25 June 1986 


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EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


THomas D. EICHLIN, Retiring Editor 


Associate Editors: 
BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON, THEODORE D. SARGENT 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


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CONTENTS 


BIRD PREDATION ON LEPIDOPTERA AND THE RELIABILITY OF 
BEAK-MARKS IN DETERMINING PREDATION PRESSURE. Mark 
K. Wourms ¢+ Fred E. Wasserman ....... Eee 


A NEW SPECIES OF CLEARWING MOTH, CARMENTA LAURELAE 
(SESIIDAE), FROM FLORIDA. Larry N. Brown, Thomas D. 
Eichlin & Wendell Snow 0... 

THE Host PLANT, ERYTHROXYLUM (ERYTHROXYLACEAE), OF 
AGRIAS (NYMPHALIDAE). Thomas S. RQy c.cccccccccceeceeennenee 

EGG DISPERSION PATTERNS AND EGG AVOIDANCE BEHAVIOR IN 
THE BUTTERFLY PIERIS SISYMBRII BDV. (PIERIDAE). Tim- 
othy A. Kellogg 0000) 0 i 

THE FOOD PLANTS OF JALMENUS DAEMELI SEMPER (LYCAENIDAE) 
WITH NOTES ON OTHER BUTTERFLIES AND ACACIA FOOD 


PLANTS. | T. J. Hawkeswood: 2 
PREDATION ON CATOCALA MOTHS (NOCTUIDAE). Auburn E. 
Brower i NaS ie 
AN ANNOTATED LIST OF THE BUTTERFLIES AND SKIPPERS OF 
LAWRENCE COUNTY, OHIO. John V. Calhoun 
SKIPPERS: POLLINATORS OR NECTAR THIEVES? B. Adrienne B. 
Venables + Edward M. Barrows ___......... EES 
PERMANENT TRAPS FOR MONITORING BUTTERFLY MIGRATION: 
TESTS IN FLORIDA, 1979-84. Thomas J. Walker .00000.. 


ADULT NOCTUIDAE FEEDING ON APHID HONEYDEW AND A 
DISCUSSION OF HONEYDEW FEEDING BY ADULT LEPI- 
DOPTERA. James B. Johnson & Michael P. Stafford ......... 


GENERAL NOTES 


Notes on the Parasitism of Rothschildia sp. Pupae (Saturniidae) in Guanacaste 
Province, Costa Riea. Allen ‘M. Young 00000. 


An Abberation of Icaricia acmon lutzi (Lycaenidae). K. D. Cannon ..... 


Natural History Notes for Aellopos ceculus (Cramer) (Sphingidae) in North- 
eastern Costa Rica.) Allen M. Young i. 


Misdirected Monarch Mating Behavior (Danaidae: Danaus plexippus) or No- 


blesse Oblige?’. Dawe Winter i200 eu 
A New Food Plant Record for Atalopedes campestris (Boisduval) (Hesperi- 
idae): Charles 'T); Bryson se 
Specimens of Callophrys rubi L. (Lycaenidae) from Figi—Transplanted Col- 
ony or One-Time Occurrence? Kurt JOR MSO oocciccceccceccceccsneeesneeccenessnnceeeeeemnnee 
Ecological Observations on Apodemia phyciodoides Barnes & Benjamin 
(Riodinidae), ° Michael]. Smith) (0 ed 
Population Outbreak of Pandora Moths (Coloradia pandora Blake) in the 
Mammoth Lakes Area, Califormia. Kelly Richer ....ccc:cs:cccecocssssssccsecsssssneeeeeeee 
Puddling by Single Male and Female Tiger Swallowtails, Papilio glaucus L. 
(Papilionidae). Theresa A. Berger & Robert C. Lederhouse -...cccccccccc 
Flower Visitation Records for Snout Butterflies (Libytheidae). Oakley Shields 
Male Determined Mating Duration in Butterflies? Per-Olof Wickman ......... 
Satyrium auretorum auretorum (Boisduval): A New Species for Oregon (Ly- 
caenidae); « Georpe 'T: Austin tice eae Oe 


INDEX TO VOLUME 39 cost pee Ne Oh a mia a lee a 


239 


262 





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